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Full text of "Phytoneuron"

I Spermo/ep/s (Ap'iaceae). Phytoneuron 2012-87: 1^9. 



TAXONOMY OF APIASTRUM, AMMOSELINUM, AND SPERMOLEPIS (APIACEAE) 

Guy L. Nesom 

2925 Hartwood Drive 
Fort Worth, Texas 76109 
guyTtesom@sbcglobal.net 

ABSTRACT 

A taxonomic summary is given for the three closely related and primarily North American 
genera Apiastrum (1 species), Ammoselinum (3 species), and Spermolepis (11 species). Ap iastru m 
includes the single species A. angustifolium, which occurs in California, Baja California, and Baja 
California Sur. Ammoselinum includes A. rosengurtii (endemic to Uruguay) and the North American 
A, butleri and A, popei, Spermolepis includes one species endemic to the Hawaiian Islands — S. 
hawaiiensis, one species endemic to Argentina — S. castellanosii, and nine species native to North 
America: S. echinata, S. mermis, Spermolepis organensis Nesom, sp. nov. (Dona Ana Co., New 
Mexico), Spermolepis laevis Nesom, sp. nov. (central Texas to south-central Oklahoma), S. 
divaricata, Spermolepis (Leptocaulis) diffusa (Nutt. ex DC.) Nesom, comb, nov., Spermolepis 
(Ammoselinum) gigantea (Coulter & Rose) Nesom, comb, nov., Spermolepis lateriflora Nesom, 
sp. nov. (California, Arizona, New Mexico, Texas, Chihuahua, and Sonora), and Spermolepis 
infernensis Nesom, sp. nov. (San Diego Co., California). Species descriptions and keys to the 
genera and species are provided and a discussion of inflorescence architecture points out distinctions 
within and among the genera; most of the species are illustrated by photos of specimens. An epitype 
is designated for Spermolepis hawaiiensis; leetotypcs are designated for Apiastrum latifolium (a 
synonym of Apiastrum angustifolium), Ammoselinum popei. Ammoselinum castellanosii 
Ammoselinum sect. Hesperoselinurn, the genus Leptocaulis (a synonym of Spermolepis), and 
Spermolepis (Leptocaulis) diffusa, 

KEY WORDS: Apiastrum, Ammoselinum, Spermolepis, Apiaceae, inflorescence architecture 



The genera Apiastrum, Ammoselinum, and Spermolepis are very similar among themselves 
and are all placed in tribe Selineae (Downie et al. 2010) of subfamily Apioideae. The species are 
primarily North American but Ammoselinum and Spermolepis each include a South American species 
and Spermolepis hawaiiensis is endemic to Hawaii. The monospecific Apiastrum is restricted to 
Pacific coastal region of Mexico (Baja California and Baja California Sur) and California. Plants of 
Ammoselinum and Spermolepis are annuals with narrow, characteristically linear to filiform leaf 
segments, narrow involucel bractlets but lacking an involucre, white petals with straight apices, 
laterally compressed fruits, narrowly conical stylop odium, and styles absent, the sigmas divergent 
Apiastrum is similar but apparently is specialized in its lack of sepals, stylopodium, and involucel 
bractlets and its branching-inflorescence architecture. Loss of peduncles has occurred in some 
species of all three genera. 

1. Medial and distal leaves appearing opposite; schizocarps dqiressed-ovoid, 1-1.5 mm; sepals 

absent; stylopodium obsolete, styles filiform; involucel bractlets absent Apiastrum 

1. Leaves alternate; schizocarps broadly ovoid to ovoid-oblong, urceolate- ovoid, or urceolate- oh long, 
1.5-5 mm; sepals small but present; stylopodium present, styles obsolete; involucel bractlets present. 

2. Schizocarp ovoid-oblong to urceolate-ovoid or broadly ellipsoid, ribs sparsely to densely 

scaberulous with single-celled papilla-like projections Ammoselinum 

2. Schizocarp broadly ovoid to ellipsoid or elliptic-ovoid, ribs and intervals variously hairy or at 
least tuberculate Spermolepis 



Nesom: Taxonomy of Apiastrum, Ammoselmum, and Spermolepis 



In maintaining Ammoselinum and Spermolepis as separate genera in the present manner, the 
definition of Ammoselinum is narrowed to only the three species with mericarps scabrous-ribbed, 
otherwise glabrous, and with corky-expanded, appendage-like lateral ribs (Fig 1). Those species 
without expanded lateral ribs and with hairs or at least tubercles on both the ribs and intervals are 
referred to Spermolepis. 

Considerable variation in fruit shape and vestiture also exists among the species of 
Spermolepis as accepted here (Fig. la-q). Spermolepis gigantea and S. castellanosii appear to be 
distinct as a pair on the basis of the relatively elongate fruits (compared to other species of 
Spermolepis), but S. gigantea is unique in its hispid-hirsutulous fruit vestiture with long, sharp- 
pointed hairs without tuberculate bases; hairs of S. castellanosii are similar to those of S. infemensis. 
Spermolepis divaricata and S. diffusa are distinct as a pair on the basis of their short-ellipsoid fruits 
with tiny upcurved hairs; they also are distinct from the rest of the genus in their relatively smooth 
epidermis (vs. minutely "bubbly'' in the others). Spermolepis echinata, S. hawaiiensis, S. lateriflora, 
S. infemensis, and S. inermis are similar in their broadly ovoid fruits with multicellular, tuberculate 
trichome bases and the first three species are echinate-brrstly with apically hooked hairs. 




Figure la-u. Mature or near-mature fruits of Apiastrum, Ai 
but not exactly the same scale (see descriptions for measure 
Ammoselinum pope i, (a) Ammoselinum rosengurtii, (d)Amj 



loselinum, and Spermolepis. At approximately 
2nts). {a) Apiastrum angustifolium, (b) 
selinum butleri. 



i: Taxonomy of Apiastrum, Ammoselinum, and Spermolepis 3 




Figure le, f, g, h. (e) Spermolepis echinata, (f) S. lateriflora, (g, h) variants ofS. ham 



n: Taxonomy of Apiastrum, Ammoselinum, and Spermolepis 4 




e ] i, j, k, 1, m, n. (i) Spermolepis echi 

text (j) S. echinata x 5. inermisf?, Cory 48' 
■is , (m) S. inermis, (n) 5. /aevw. 



Nesom: Taxonomy of Apiastrum, Ammoselinum, and Spermolepis 5 




Figure lo, p, q, r, s, t. (o) Spermolepis laevis variant with slightly rugulate ribs and intervals - Wolff '2102, (p) S. 
laevis variant with a few tubercles, some with short, blunt-tipped hairs - Whitehouse 18439, (q) S. divaricata, (r) 
S. diffusa, (b, t) S. castellanosii 



Nesom: Taxonomy of fyfatfrun, Ammosefaium, aid Spermdepe 



. . t 





4**J& ■ 






u 



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Figure lu. ifyxrmolepis gigantea (at much larger scale than the other fruit photos). 



Among the genera of tribe Selineae in the account by Downie et al. (2010), Oligzclaaus 
Chodat & Wilczek is the only other American genus in the tribe outside of the "Arracacia Clade" and 
the "Perennial Endemic Norm American Clade" — except for Ammoselimim and Spermolepis, all 
other genera are Eurasian. Oligzclaatis includes only the single species O. pala^micas (Speg.) 
Perez- M or. (synonym = O. anehmts Chodat & Wilczek), which apparently is restricted to Argentina. 
Mainly because of its dorsally compressed fruits (flattened parallel with the plane of the commissure) 
with numerous oil tubes on the broad commissural face, M ami as and Constance (1950) eliminated 
Oligzclaatis as a possible congener or even close relative when considering the generic placement of 
their new species Ammoselimim rosengirtii (see illustrations of O. amtmis in Chodat and Wilczek 
1902, pp. 527—528, vs. lateral compression in the Apiastnim-Ammoselimim-Spermolepis group, 
flattened perpendicular to the plane of the commissure). 

The base chromosome number of the Apiastrum, Ammoselimim, and Spermolepis group 
appears to be x = 11, as it appears in all three genera, with reductions to x = 10 and x = 8. Among the 
uncinate-bristly species of Spermolepis, three dysploid levels exist: 2n = 22 (S hcrwaiiensis), 2n = 20 
(S ecHnald), and 2n = 16 (S lateriflora). If the uncinate-bristly species represent a single clade, men 
the dysploid changes appear to be more indicative of individual speciation events that consistent 



: Taxonomy of Apiastrum, Ammoselinum, and Spermolepis 



indicators of cladistic relations hips. Interesting research remains to be done with regard to 
chromosome numbers (see comments following S. divaricatd). 

A close ifclduonsh,} at u.»ij ihe 1 3 species of Apiastrum, Ammoselinum, and Spermolepis is 
suggested by morphological similarities as well as their general geographic coherence in a broad 
region (North America, South America) where other potentially related species apparently do not 
exist. It would not be indefensible to treat all 13 species within a single genus. 

Branching pattern and umbel architecture 

In all species of Ammoselinum except one and in two species of Spermolepis, umbels are 
borne on ebracteate peduncles mat appear to originate only at leaf axils (Fig. 2A). Growth is 
indeterminate, as upward vegetative growth is continued even at the distalmost node. In these plants, 
however, the peduncle appears to be the extension of the primary ("pr") stem axis. Continued upward 
stem growth and production of additional umbels continues from growth of the axillary ("ax") bud. 
In this interpretation, the umbels actually are produced as terminal structures, rather than axillary ones. 

In Ammoselinum butleri, Spermolepis lateriflora, and S. infernensis, the peduncle is absent 
(presumably suppressed) and the umbellet rays appear to arise from the leaf axils (Fig. 2C). The 
axillary bud in these plants apparently is suppressed ;it the distalmost node, so that growth may be 
characterized as determinate. Peduncle suppression is complete in A. butleri but in S. lateriflora, 
apparently over its whole geographic range, some plants produce pedunculate umbels from nodes 
below the distalmost (see Figures 7 and 8 and examples cited below, under the species). 

In Spermolepis divaricata, S. diffusa, and S. inermis, the primary axis at each node forms a 
peduncle and compound umbel, but on the branch arising from the distalmost axillary bud, both the 
terminal leaf and the axillary bud are suppressed (Fig. 2B). In effect, each branch terminates in two 
compound umbels and growth may be characterized as determinate. 

In Apiastrum angustiflolium, leaves appear to be opposite and two branches and two sessile 
(compound) umbels arise at each node (Fig. 2D). In the interpretation hers, this arises from (a) 
suppression of the peduncles, (b) complete foreshortening of the distal internode that would constitute 
the axillary axis, and (c) duplication of the axillary bud in order that growth continues from two 
upward branches. Axillary buds apparently are suppressed at the distalmost node and growth may be 
characterized as determinate. 



Nesom: Taxonomy of Apiastrum, Ammoselinum, and 




Figure 2. Variation in inflorescence architecture in Ammoselinum., Apiastrum, and Spermolepis. A. 
Pedunailaie-axillary-in determinate: Ammoselinum popei, Ammoselinum rosengurtii, Spermolepis eciunaM, S. 
hawaiiensis, & castelkmosii. B. Pedunculate- axillary-determinate: Spe}tiv:Aepi.s divaricata. S. diffusa., S 
inermis, S. laevis, S. orgaiiensis, S. gigantea. C. Sessile-axil lary-deteraiin ate: Ammoselinum butleri, 
Spermolepis lateriflora, S. infernensis. D. Sessile-dichotomous- determinate: Apiastrum angustifoUum. "ax" = 
axillary brand], arising from the axillary bud. "pr" = primary branch, continuing from the main branch from 
below. In determinate arrangements, rise axillary bud is suppressed at the distaimost node. 



: Taxonomy of Apiastrum, Ammosdinum, and Spermolepis 



APIASTRUM Nutt. ex Torr. & A. Gray, Fl. N. Amer. 1: 643. 1840. TYPE: Apiastrum angustifolium 
Nutt. ex Torr. & A. Gray 

Herbs, annual, with odor, 0.4-5 dm, glabrous; taproot slender. Leaves appearing opposite to 
subopposite medially and distally; basal and cauline 3-4 ternately compound; blades broadly ovate to 
ovate in outline, herbaceous; leaflets divided, ultimate divisions linear to narrowly oblong, margins 
entire; petioles scarious margined at base, distal petioles foreshortened and scarious-margined along 
whole length. Umbels compound, loosely convex, axillary, sessile (rays appearing to arise from leaf 
axils), peripheral flowers not different; involucral bracts absent; involucel bractlets absent. Pedicels 
present. Flowers bisexual; sqjals absent; petals white, margins, entire, apices slightly inflexed; 
stylopodium depressed-reduced, nearly obsolete; styles evident, filiform, 0.2 mm, arching-divergent. 
Schizocarps depressed-ovoid [mericarps reniform], laterally compressed, not beaked, splitting, ribs 3. 
barely raised, mostly delimited by line of papillae, oil tubes 1 per interval, filiform not filling the 
interval, surface shallowly tuberculate, otherwise glabrous; carpophore bifid the whole length. Base 
chromosome number, x = 11. 

1. Apiastrum angustifolium Nutt. ex Torr. & A. Gray, Fl. N. Amer. 1(4): 644. 1840. TYPE: USA. 
California. [San Diego Co.:] St. Diego, N Cal, April, T. Nuttall s.n. (probable holotype: GH 
00075076; isotypes: K digital image!, PH 01015720 digital image!). 

Apiastrum angustifolium var. tenellum Nutt. ex Torr. & A Gray, Fl. N. Amer. 1(4): 644. 1840. TYPE: 
MEXICO. Baja California. Cerro [Cedros] Island, Mar 1889, E. Palmer s.n. (probable 
holotype: PH 743994 digital image!). 

Torrey and Gray did not indicate that they saw a collection; their concept of the taxon was 
from Nuttall's manuscript. The plant in the PH collection matches Torrey and Gray's brief description 
("stem dichotomous from the base; leaves less divided; rays of the umbel very slender; umbellets 1-2- 
flowered; seed morerugulose"). 

Apiastrum latifolium Nutt. ex Torr. & A Gray, Fl. N. Amer. 1(4): 644. 1840. LECTOTYPE 
(designated here): USA. California. [Santa Barbara Co.:] "St. Barbara, N Cal." [on PH 
sheet], no other collection data, T. Nuttall s.n. (GH 00075075; isolectotype: PH 01044838 
digital image!). Torrey and Gray cited "Nuttall! Douglas!" 

Helosciadium leptophyllum var. ? latifolium Hook. & Arn., Bot. Beechey Voy., 347. 1838. No 
collection was cited (pp. 347-348). The protologue gave only this: "The specimens are only 
in young fruit, and the segments of the upper leaves are considerably broader than in any 
form we have yet seen, while even the lower ones are broader than in H. laciniatum, DC, 
which we consider a mere variety of this species." As synonym of Apiastrum fide Mathias 
and Constance (1945). 

Stems 4-50 cm. Leaves: blades 1-5 cm, ultmutw segments 5-25 mm; petioles 20-40 mm. 
Peduncles absent. Umbels axillary only: involucre bracts absent; involucel bractlets absent; fruiting 
rays 2-5 per node (2 umbels per node), (0-)7-25(-50) mm (central umbellet sessile); umbellets 3-7 
flowered; fruiting pedicels (0-)2-8(-15) mm (central l-2(-3) flow^ers sessile), unequal, spreading. 
Schizocarps 1-1.5 mm. In = 22 (San Diego Co. - 2 counts, Bell & Constance 1957; Baja California, 
Constance et al. 1976). Map 1. Figure 3. 

Flowering Mar-Apr, Chaparral, coastal sage scrub, blue oak savanna, rock outcrops, granite 
slopes, shale slopes, serpentine soil, steep slopes, recently burned areas, grassy openings, roadsides; 
0-400(-1500) m. Calif.; Mexico (Baja California. Baja California Sur). 



n: Taxonomy of Apiastrum, Ammoselinum, and Spermolepis \Q 



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Map 1. Distribution of Apiastrum ang/jstifohm 



: Taxonomy of Apiastrum, Ammosdinum, and Spermolepis 



AMMOSELINUM Torrey & A Gray, Pacif. Railr. Rep. 2(4): 165. 1855 [1857]. TYPE: 

Ammosdinum popei Torrey & A. Gray 

Herbs, annual, odorless or "faintly Pasttnaca-scented" in A. gtganteum, 0.4-3.5 dm, glabrous 
but stem ridges distally scaberulous; taprooted. Leaves all alternate; basal 3-ternately compound, 
cauline 2-3-ternately compound; blades broadly ovate to obovate, herbaceous; leaflets lobed or 
divided, leaflets and ultimate divisions linear to oblanceolate or subspatulate, margins entire; petioles 
scarious margined at base, distal petioles foreshortened and scarious-margined along whole length. 
Umbels compound, loosely convex, axillary, pedunculate (rays borne on an ebracteate portion of 
stem) or sessile (rays appearing to arise from leaf axils), peripheral flowers not different; involucral 
bracts absent; involucel bractlets distinct, linear, entire or less commonly 2-3-fid, herbaceous. 
Pedicels present. Flowers bisexual; sqiais obsolete or greatly reduced and barely evident; petals 
white, margins entire; stylopodium narrowly conical; styles obsolete, stigmas directly atop 
stylopodium and divergent. Schizocarps ovoid-oblong to urceolate-ovoid or broadly ellipsoid, 2-5.3 
mm, laterally flattened, base shallowly rounded to truncate, apex not beaked, mericarps splitting, 
dorsal ribs 3, corky-thickened or thin, lateral corky-thickened with an appendage-like growth 
incurving over the commissure, dorsal oil ducts 1 or 3 per interval, 1 or 2 on the commissure, ribs 
scaberulous with rounded to blunt single-celled, irregular, papilla-like projections; commissure 
sulcate; carpophore bifid only distally or along the whole length. Base chromosome number, x = 1 1. 

Key to the species 

1. Umbels epedunculate (sessile, rays appearing to arise from leaf axils); schizocarps 2-2.2 mm, 

dorsal ribs acute 1. Ammoselinum butleri 

1. Umbels pedunculate (rays borne on an ebracteate portion of stem); schizocarps 3^1(-5) mm, dorsal 
ribs rounded or acute. 

2. Mature fruits tan; ribs usually corky-thickened, not thin and wing-like; North America 

2. Ammoselinum popei 

2, Mature fruits blackish; ribs thin, wing-like; South America 3. Ammoselinum rosengurtii 

1. Ammoselinum butleri (Engelm. ex S. Wats.) Coulter & Rose, Bot. Gaz. 12: 294. 1887. Apium 
butleri Engelm. ex S. Wats., Proc. Amer. Acad. Arts 21: 453. 1886. LECTOYPE (Coulter & 
Rose 1900, p. 90): USA. Texas. Harris Co.: near Houston, 29 Mar 1872, E. Hall 244 (GH; 
isolectotypes : BM digital image!, K digital image'). 

Watson cited three collections: (1) "Texas, in wet grounds near Houston, E. Hall (n. 244), 
March, 1872"; (2) "near Dallas, J. Reverchon[s.n.\ March and Aprii 1874"; and (3) "Indian territory, 
south of the Arkansas, CD. Butler [s.n], 1876." Coulter and Rose (1900) noted that "The type here 
gn-en is the first specimen cited" but they clearly referred to it as the type and to the Reverchon and 
Butler collections as "associated with it in the original description." 

Stems 4-10 cm, branching from the base. Leaves: blades oblong to oblong-ovate in outline, 
10-25 mm, ultimate divisions 1-8 mm; leaflet and bractlet margins and midrib smooth to very 
sparsely scaberulous; petioles 5-30 mm, clasping to auriculate-clasping, scarious-margined at the 
base. Inflorescence axillary, indeterminate. Peduncles absent (rays appearing to emerge from the 
leaf axils). Rays (l-)2-4, (0-)0.5-4(-7, very rarely to 15) mm. Umbellets (l-)3-5-flowered (central 
umbellet often 1-flowered); pedicels (0.5-)l-3(-4) mm (central flower sessile). Involucel bractlets 
1-4, linear, rarely 2-fid, 0.5-2 mm, without scarious margins. Schizocarps ovoid-oblong, 2-2.2 mm, 
dorsal ribs acute, sparsely papillate-scaberulous with minute, translucent, apparently 1-celled, conical, 
apically acute, papilla-like hairs, lateral ribs slightly corky-thickened, barely raised; oil tubes 1 per 
interval, filiform but usually clear!}' evident, 2 on the commissure; carpophore bifid in distal 1/4. 
Chromosome number not reported. 



: Taxonomy of Apiastrum, Ammosdinum, and Spermolepis 



Flowering Mar-Apr. Sandy and sandy clay soil, gravel piles, lawns, old fields, roadsides, 
cultivated fields, pastures, fencerows, stream and pond edges, shell banks, oak-juniper glades, 
limestone prairies; 100-300 m; Ala., Ark., Kans., La., Miss., Mo., N.C., Okla., Term., Tex. 

Almost all habitats recorded for Ammoselinum butler i have been described in some sense as 
"disturbed" — some of those in Texas (glades and prairies) apparently are undisturbed. The species 
has only recently expanded into Alabama, Mississippi, and North Carolina (Boufford 1977; Bryson 
1991; Keener 2007). 

2. Ammoselinum popei Torrey & A Gray, Pacif. Railr. Rep. 2(4): 165. 1855 [1857]. Apium popei 
(Torrey & A Gray) A. Gray, Proc. Amer. Acad. Arts 7: 343, 1868. LECTOYPE (designated 
here): USA. Texas. Headwaters of the Colorado, 13 Apr 1854, J. Pope s.n. (NY digital 
image!). At GH is another Pope specimen, labeled "Mar-Apr, Llano Estacado;" this may be a 
duplicate of the NY sheet, but it is not clear. 

The protologue has this: ''Sandy soil: Llano Estacado, and head-waters of the Colorado 
[collections by 'Captain Pope']; March and April. Mr. Wright found it in Western Texas, but he 
collected only a few specimens, and it was not distributed with his plants. Some ripe seeds that he 
collected were cultivated in the Cambridge Botanic Garden, and arrived at perfection. Dr. Parry, while 
engaged on the Mexican boundary survey under Major Emery, sent home a single flowering specimen 
of the plant, found at Eagle Pass in January, 1853. From no other sources have we received any 
specim ens of this apparently new genus. " 

Stems 8-35(-60) cm. branching from the base. Leaves: blades oblong-ovate in outline, 10- 

40 mm, ultimate divisions 2-10 mm; leaflet and bractlet margins and midrib prominently scabrous. 
petioles 5-60 mm, clasping to auriculate-elasping, scarious-margined at the base. Inflorescence 
axillary, indeterminate. Peduncles 25-75 mm. Rays (5-)6-10, (0-)6-25 mm (inner umbellet 
sessile). Umbellets 4-9 flowered; pedicels (0-)2-5 mm (inner flower sessile to subsessile). 
Involucel bractlets 1-6, linear and entire or less commonly 2-3-fid, 2-12 mm, usually scarious- 
margined at the base. Schizocarps urceola're-oblong, 3-4(-5) mm, dorsal ribs rounded, densely and 
coarsely papillate-scaberulous with translucent, multicelled, convex, apicaily rounded, papilla-like 
hairs, lateral ribs corky-thickened and raised and obscuring the commissural face; oil tubes 1 per 
interval, barely evident between the thickened ribs, commissural usually not evident; carpophore bifid 
along whole length. Chromosome number not reported. 

Flowering (Mar-)Apr-May. Sandy soil, rocky soil, rock outcrops, roadsides, pastures, lake 
edges, dunes, gypsum flats, limestone barrens, cedar glades, mesquite savannas, gypsiferous, 
calcareous, and black clay prairies; 0-800(-1300) m; Kans., N.Mex., Okla., Tenn, Tex.; Mexico 
(Coahuila, Nuevo Leon, Tamaulipas). 

The disjunct populations in central Tennessee (5 counties, see TENN 2012) are typical in 
morphology and apparently native there. 

An epetiolate leaf rarely is produced at the base of a cluster of rays. This is a consistent 
feature in the plants of Tolstead ''015 (SMU) from Taylor Co., Texas. 

Collections examined. USA New Mexico. Eddy Co. : Carlsbad Cavern Natl. Park, road to 
sewage lagoon just across E boundary of park, SW of Whites City, bajada below escarpment, xeric 
shrubland, Larrea-Gutierrezia community, ca. 3640 ft, 19 Apr 1977, Burgess 4449 (TEX); Carlsbad 
City Lake and park on the Pecos River, wet sandy loam, 20 Apr 1966, Crutchfield 1336 (LL); Lincoln 
Natl. Forest, Sitting Bull Falls, growing in lawn at picnic site near trail to falls, 4595 ft, 21 Apr 2011. 
Heil 33359A (SJC). Lea Co. : City of Hobbs, jet of US 62-180 and the hwy to Lovington, lawn of 
: store, 3670 ft, 23 May 2011, Heil & O'Kane 33424 (SJC); Hobbs, near Humble City, 



: Taxonomy of Apiastrum, Ammosdinum, and Spermolepis 



entrance to Ocotillo Golf Course, weedy sites, disturbed sites, 3700 ft, 20 Apr 2011, Heil 33347 
(SJC). Ot ero C o.: Escarpment and canyon in limestone plateau ca. 11 air mi NE of Dell City, 3.3 
road mi N and then NE from the TX/NM state line and E from hwy up canyon for 0.75 mi, on 
alluvium below limestone slopes, 3706 ft, roadside plants, 2 May 1999, Worthington 28247 (UCR, 
fide SEINET). 

Collections examined. MEXICO. Coahuila. Musquiz, 12 Apr 1936, Marsh 2097 (TEX); 
Musquiz, Apr 1938, Marsh 1132 (TEX); 56 mi S of Eagle Pass, Texas, rocky slope, 3 April 1969, 
Pinkava 15576 (ASU digital image!); Rio Grande valley, near Diaz, 700 ft, 15 Apr 1900, Pringle 
8314 (MO digital image!). Nuevo Leon. 2 mi N of Sabinas Hidalgo, sandy loam bottom, 26 Mar 
1944, Barkley 14521B (TEX); 23 mi N of Sabinas Hidalgo, limy clay hillside, 26 Mar 1944, Barkley 
14592 (TEX); 5 km N of Sabinas Hidalgo on Hwy 85, dry but verdant wash leading from apparently 
irrigated field, surrounding vegetation of Acacia-Prosopis, 3 10 m, 23 Mar 1986, Nesom 5350 (TEX); 
Mpio. Higueras, Cuesta Mamulique, ca. 40 km S of Sabinas Hidalgo on Mex 85, W side of "Libre" 
Hwy, near top of ridge of dirt road toward microwave tower, area of Acacia and other genera of 
shrubs, ca. 540 m, 26 Mar 1993, Nesom 7553 (TEX); fields near Monterrey, 1800 ft, 6 Apr 1906, 
Pringle 13747 (LL. TEX). Tamaulipas. 10 mi S of Nuevo Laredo, dry flat, 26 Feb 1944, Barkley 
14322 (TEX); 3 mi SW of El Canelo at lat. 25° 09' on the Matamoros-Victoria hwy, clay roadside 
ditch, 50 ft, 9 Feb 1960, Johnston 5077C (TEX). 

3. Ammoselinum rosengurtii Mathias & Constance, Bull. Torrey Bot. Club 77: 133, fig. 1. 1950. 
TYPE: URUGUAY. Depto. Florida, Estancia Rincon de Santa Elena, Picada Castro, Arroyo 
Mansavillagra, 8 Nov 1946, B. Rosengurtt Gallinal 5753 (holotype: UC! digital image!; 
isotype: LA). 

Stems ca. 4-8(-10) cm tall, simple or few-branched mostly at 1 or 2 nodes above the base; 
"from a Daucus-scsnted taproot" (Rosengurtt in 1969) Leaves: blades broadly ovate in outline 25- 
40 mm, ultimate divisions 4-8 mm, scaberulous on margins and nerves; petioles 7-10 mm, scarious- 
margined along whole length. Peduncles 20-50 mm. Rays 2-5, 1-20 mm (inner 1-2 umbellets 
short-pedicellate). Umbellets (l-)3-5 -flowered, pedicels 2-7(-10) mm (central flower short- 
pedicellate). Involucel bractlets 3-4. linear-lanceolate, entire, 1.5-7 mm, unequal, without scarious 
margins. Schizocarps ellipsoid, attenuate toward the apex, 3.2-5.3 mm, densely scabrous on the 
angles with thick, sharp-pointed, pustulate-based hairs, dorsal ribs 3, rounded, lateral ribs; oil tubes 3 
per interval, 2 on the commissural face; carpophore bifid along whole length. In = 44 (from 
Rosengurtt s.n., 1969, see citation below; Constance et al. 1976). Figures 4, 5. 

Flowering Oct-Dec. Habitat?; ca. 100-200 m; South America (Uruguay). 

Additional collection examined. Uruguay. Depto. Rocha, Sta. Teresa, Dec 1969, Rosengurtt 
s.n. (TEX), chromosome voucher cultivated in Univ. of California Botanical Garden C-1721. Also 
see photos on Flickr (Gonzalez 2010). 

Ammoselinum rosengurtii apparently is known only from Uruguay (Mathias & Constance 
1950); the protologue cited collections from provs. Artigas. Cerro Largo, Florida, Paysandu, Rio 
Negro, Salto, and Soriana. Photos by Andres Gonzalez were taken "al margen del Rio Negro cerca al 
dique de la Represa Constitucion. departamento de Rio Negro." 

Ammoselinum rosengurtii is similar to A. popei especially in its papillate-scabrous vestiture, 
urceolate-oblong fruits with expanded lateral ribs, and carpophore divided along the whole length. It 
differs from A popei in its dark fruits, thin-winged fruit ribs, and three oil tubes per dorsal interval. 



: Taxonomy of Apiastrum, Ammosdinum, and Spermolepis 



SPERMOLEPIS Raf, Neogenyton, 2. 1825. TYPE: Spermolepis divaricate! (Walt.) Raf. ex Seringe 
In the protologue, Rafinesque noted "Type, a plant put in four genera already! Sison 
pusillum, Mx. Daucus divaricates, Walt. Ammi do [divarieatum] Pers and Ligusticum 
pusillura Pers ! " The nomenclatural combination iji Spermolepis for Daucus divaricatus Walt, 
was made by Seringe in 1830, attributed by him to Rafinesque. 

Leptocaulis Nutt. ex DC, Coll. Mem. 39, plate 10. 1829. LECTOTYPE (designated here): Leptocaulis 
divaricatus (Walt.) DC. De Candolle's discussion included, in various permutations, 
Spermolepis divaricata, S. inermis, S. echinata, and various synonyms. 

Babiron Raf, New Fl. 4: 23. 1838. TYPE: Babiron divarieatum (Walt.) Raf. = Spermolepis 
divaricata Rafinesque included Babiron pusillum, B. divarieatum. and B. dichotomum — all 
are synonyms of Spermolepis divaricata. 

Lepisperma Raf, Act. Soc, Linn. Bordeaux 6: 268. 1834. As synonym of Spermolepis fide Mathias 
and Constance 1945). 

Ammoselinum sect. Hesperoselinum Munz & Johnston, Bull. Torrey Bot. Club 52: 224. 1925. 
LECTOTYPE (designated here): Ammoselinum giganteum Coulter & Rose 
After identifying their new species Ammoselinum occidentals Munz & Johnston as congeneric with A. 
giganteum, the authors placed the two species together in a separate section. No type was designated. 

Annual herbs, slender taprooted, glabrous, not aromatic or sometimes (S. lateriflora) with a 
"carrot" odor. Stems erect, 5-80 cm, simple or few-branched from basal to medial nodes. Leaves all 
alternate; basal and cauline or mostly cauline, 3-pinnately compound; blades ovate to oblong to 
oblong-ovate in outline; leaflets filiform to linear or narrowly oblong, margins entire to weakly 
scaberulous; proximal petioles scarious margined at base, distal petioles becoming much 
foreshortened and scarious-margined along whole length. Umbels compound, irregularly and loosely 
convex, termin.il and axillary or axillary only, pedunculate (rays borne on an ebracteate portion of 
stem) or sessile (rays appearing to arise from leaf axils), peripheral flowers not different; involucre 
bracts absent or (S. hawaiiensis) sometimes present; involucel bractlets distinct, filiform or linear to 
linear-lanceolate, herbaceous, without scarious margins. Pedicels present or reduced to obsolete. 
Flowers bisexual; sqials obsolete or essentially absent petals ^Litc oblong or elliptic to ovate, apex 
not infiexed, margins entire; stylopodium conical; styles obsolete, stigmas directly atop stylopodium 
and divergent. Schizocarps broadly ovoid to ellipsoid or elliptic-ovoid, sometimes slightly beaked, 
1.5-2(-4) mm, flattened laterally and slightly constricted at the commissure, mericaips splitting, ribs 
[number], filiform, oil tubes l(-3) pel' interval, 2 on the commissure, surface (a) echinate-bristly with 
apically hooked hairs arising from rounded, multicellular bases, (b) bristly with apically straight hairs 
arising from rounded, multicellular bases, (c) tuberculate-roughened by rounded, multicellular 
projections, (tuberculae and multicellular hair bases apparently homologous), or (d) scabrous with 
minute, upcurved hairs not arising from a tubereulate base; commissural face sulcate; carpophore 
bifid in distal 1/4-1/3. Base chromosome number = 11? (n = 8, 10, 11, 19, 32). 

KEY TO THE SPECIES 

1. Schizocarps densely echinate-bristly with sharp-pointed, apically hooked hairs. 

2. Distal umbels sessile, without a peduncle, proximal umbels sometimes pedunculate; In = 16 

1. Spermolepis lateriflora 

2. All umbels distinctly pedunculate. 

3. Schizocarps 1.5-2 mm; 2n = 20 2. Spermolepis echinata 

3. Schizocarps 3^1 mm; 2/3 = 22 3. Spermolepis hawaiiensis 

1. Schizocarps with apically straight hairs or lacking hairs. 



: Taxonomy of Apiastrum, Ammoselinum, and Spermolepis 



4. Distal umbels sessile, without a peduncle, proximal umbels sometimes pedunculate; fruit surface 
tuberculate on ribs and intervals, some tubercles with apically straight hairs, some without hairs 

4. Spermolepis infernensis 

4. All or most umbels distinctly pedunculate; fruit surfaces variable in\ 



5. Schizocarps 3-5 mm, narrowly elliptic- ovate to oblong-ellipsoid or ovoid-oblong, surface 
hispid-hirsutulous. 

6. Hairs blunt-tipped, arising from a tuberculate base; South America 
10. Spermolepis castellanosii 

6. Hairs sharp -pointed, arising from a non-tub erculate base; North America 
11. Spermolepis gigantea 

5. Schizocarps 1.2-2 mm, mostly broadly ovate to broadly ellptic, surface either completely 
smooth, tuberculate but hairless, tuberculate with a few hairs, or sparsely to densely scabrous 
with minute, upcurved or upturned hairs. 

7. Fruit surface ribbed but otherwise completely smooth, lacking even tubercles 

6. Spermolepis laevis 

7. Fruit surface either tuberculate (with or without straight, erect hairs) or with minute, 
upcurved or upturned hairs. 

8. Fruit surface tuberculate with multicellular trichome bases, but totally lacking hairs 

9. Tubercles irregularly scattered, some with short erect hairs; peduncles 0.9-3,5 cm 
7. Spermolepis organensis 

9. Tubercles densely arranged, without hairs; peduncles 2-7 cm 
5. Spermolepis inermis 

8. Schizocarps sparsely to densely scabrous with minute, 1-celied, upcurved or upturned 
hairs, the hairs not arising from multicellular tubercles. 

10. Pedicels (0-)2-9 mm; rays 5-17 mm; central 1-2 flowers of each umbellet sessile to 

subsessile 8. Spermolepis divarieata 

10. Pedicels (8-) 14—32 mm; rays 15-33 mm; all flowers of each umbellet with subequal 
pedicels, none sessile or subsessile 9. Spermolepis diffusa 



1. Spermolepis lateriflora G.L. Nesom, sp. nov. TYPE: USA. Arizona. Pima Co.: Rillito Valley, 
Tucson, 29 Apr 1905, J.J. Thornber 5241 (holotype: TEX!; isotypes: ARIZ!, RSA!, UC!). 

Similar to Spermolepis echinata (and previously identified as that species) in its fruits densely 
echinate-bristly with uncinate hairs but distinct in its cpedunculate (sessile) umbels. 

Stems 5-35 cm. Leaves: blades broadly ovate in outline, mostly 1-5 cm, finely temately 
dissected, ultimate segments linear to oblong, 4-12 mm; petioles 1-3 cm. Peduncles absent or 20-70 
mm. Umbels usually axillary only, usually sessile at all nodes, alway sessile at distal nodes, 
occasionally pedunculate below the distalmost; involucre bracts absent; involucel bractlets 2-4, linear, 
1-2 mm; fruiting rays 4-5 per node, (0— )1— 14 mm (central 1-2 umbellets sessile to subsessile), 
unequal, spreading; umbellets 3-8-flowered; fruiting pedicels 1-6 mm (central 1-2 flowers 
subsessile). Schizocarps 2-2.2 mm, densely echinate-bristly with apically hooked hairs; oil tubes 1 




Map 2. Distribution of Spermokpis lateriflora. 

Additional collections examined. USA. California. [Alameda Co. :] Oakland Hills, May 
1877, [J.G. Lemmon?] s.n. (UC!). Los Angeles Co. : Verdugo Mountains. Tuna Canyon, shady moist 
banks, 1100 ft, Apr 1930, MacFadden 2441 (MACF digital image!). San Diego Co. : due S of 



: Taxonomy of Apiastrum, Ammosdinum, and Spermolepis 



BoiTego Valley, rocky terrain on alluvial slopes. 1250 ft, 30 Mar 1952, Bacigalupi &Macbride 3570- 
A (VC\); lower Box Canyon, 29 Apr 1941, Gander 9194 (SD fide Calif. Consortium); near base of 
hills S of Vallecito Stage'Sta., 30 Apr 1941, Gander 9260 (SD digital image!). Other map points are 
from specimens studied from ARIZ, BRIT-SMU-VDB, NMC, SRSC, TEX-LL, and UC-JEPS. 

Munz and Keck (1959) identified the California plants as Ammoselinum giganteum (with A 
occidentale in synonymy) but, following their identification/annotation as Spermolepis echinata in 
1948 by Constance, they have been identified as S. echinata in iterations of the Jepson Manual 
(Constance 1993; Constance & Wetherwax 2012). 

All of the California collections of Spermolepis lateriflora are sq^arated from the main range 
of the species and all were made near urban areas (Oakland, Los Angeles, San Diego). These 
occurrences perhaps resulted from inadvertent recent dispersal of the echinate-bristly fruits from 
Arizona. CalFlora (2012) shows a number of other records (identified as S. echinata, apparently not 
vouchered) from the southwest region of the Anza-Borrego Desert State Wilderness Park (about 50- 
60 miles east of San Diego) and these may be other occurrences of S. lateriflora. The Vallecito Stage 
Station (Gander 9260) is on the southern border of the Anza-Borrego park. The type locality of S. 
infernensis (Hellhole Canyon Preserve, as described below), however, is in the close vicinity 7 and the 
identity of these unvouchered records needs to be verified. 

Additional collections examined. USA. Texas. El Paso Co. : E slope of Franklin Ml, N of El 
Paso, off War Road, bajada, 1 May 1970, Gorrell 38548 (LL); Franklin Mts. on trail to Cottonwood 
Springs, E of Canutillo, on bajada, 12 May 1959, Correll & Johnston 21824 (LL); 0.5 mi W of 
intersection of Hwy 54 and transmontane hwy, along roadside, red sandy soils, I Apr 1998. Turner 
98-29 (TEX); ca. 20 mi E of El Paso, 4000 ft, 5 Apr 1958, Warnock & Johnston 16215 (SRSC); E 
slopes of Franklin Mountains, gravelly granitic soil, 20 Apr 1975, Warnock 23985 (SRSC); Franklin 
Mts., below (E) Fusselman Canyon flood control dam, 4450 ft, mixed alluvium, 1 Apr 1979, 
Worthington 4219 (TEX); Hueco Mts., Hueco Tanks State Park, N end of North Mountain in small 
canyon, 29 Apr 1979, Worthington 4407 (BRIT); Franklin Mts., E side of mts., 0.2 mi N jet Trans- 
Mountain Rd and War Rd, granite soil alluvial bajada, desert shrub, 4000 ft, 9 Apr 1982, Worthington 
8071 (BRIT). 

Representative collections examined (see SEINET 2012 for many others from Arizona and 
New Mexico). USA. Arizona. Cochise Co. : 19 mi NE of Douglas, desert grassland, 4000 ft, 17 Apr 
1940, Benson 10298 (ARIZ); Mule Mts., waste spots, S-facing slopes. 25 Apr 1952, Goodding 47-52 
(ARIZ). Gila Co. : Tonto National Monument, along power line, 26 Jul 2001, 740 meters, West 1099 
(ARIZ). Graham Co. : Tanque, 1200 m, 8 May 1924, Eggleston 19889 (ARIZ). M aricopa Co. : Sand 
Tank Mts., 48 km SE of Gila Bend, small drainage, Sonoran desert scrub, 30 Apr 2003, Baker 15323 
(ARIZ); Sonoran Desert Natl. Monument, Sand Tank Mts., summit of small peak 730 m NW of 
Bender Spring, 17 Apr 2001, Felger 01-362 (ARIZ). Moha ve Co. : 7 mi S of Yucca, along sandy 
wash in Joshua tree-creosote bush area, 2300 ft, 1 2 Apr 1947, Gould & Darrow 4318 (ARIZ); Yucca, 
14 May 1884, Jones s.n. (POM). Pima Co. : Fresnal Canyon, 23 Apr 1932, Harrison et al. 8600 
(ARIZ); Santa Catalina Mts., Lower Sabino Canyon, moist sand along stream, 2800 ft, 8 Apr 1946, 
Gould 3488 (ARIZ); Coronado Natl. Forest, S on State Hwy 83, ca. 13 mi from Interstate 10, open 
pastures. 1400 m, 19 Apr 1998, Schmidt &Merello 2670 (BRIT, RSA, DNA sample). Pinal Co. : Gila 
River bottom near Sacaton, 23 Feb 1926, Porter et al. 863 (ARIZ); Oracle, 12 May 1905, Thornber 
s.n. (ARIZ). Santa Cruz Co. : Nature Conservancy Patagonia-Sonoita Creek Sanctuary, SW of 
Patagonia, ca. 4000 ft, 30 Apr 1977, Fay 241 (ARIZ); 2 mi E of Nogales of Patagonia road, gravel 
along roadcut, 3800 ft, 9 May 1945, Gould & Pultz 3093 (ARIZ). Yavapai Co. : Bald Hill, mouth of 
West Cochise Stronghold, flats of mouth of canyon, 11 Apr I960, Goodding 49-60 (ARIZ); Congress 
Junction, 3000 ft, 4 May 1903, Jones s.n. (POM). New Mexico. Dona A na Co.: White Sands Missile 



: Taxonomy of Apiastrum, Ammosdinum, and Spermolepis 



Range, Gate 4. up road ca. 1 rni "Anemone Ridge.' 1 ca. 2 mi N of US 70, just E of San Augustin Pass, 
Chihuahuan Desert Scrub community, monzanite boulders, 4500 ft, 22 Mar 2010, Heil & Anderson 
32092 (SJC); W base of Organ Mtns, at the mouth of Dripping Springs Canyon, 1 1 mi E of northern 
Las Graces, about 1/4 mi W of the rains of an abandoned resort hotel [Van Patten's], plants very 
common in a flat, unshaded area at the mouth of a rocky arroyo, 6000 ft, with Erigeron nudiflorus, 
Cercocarpus, Fallugia, Celt's and Opuntia, 25 April 1982, Ward & Soreng 82-009 (NMC); in the 
Mesilla Valley; near Las Cruces, ca. 3850 ft, 8 April 1907, Wooton & Standley s.n. (NMC). Grant 
Ca: 1 mi S of Gila, gravelly mesa slope, 21 May 1935. Maguire et al. 11530 (ARIZ); 1 mi S of Red 
Rock, gravelly mesa top, assoc. Prosopis and Opuntia, 21 May 1935. Maguire et al. 11530 (UC). 
Hidalgo Co. : State Hwy 92 to Virden, ca. 2.5 mi N of US 70, roadside alluvial soils, desert grassland 
with scattered creosote, 4085 ft, 20 Mar 2010, Heil 32041 (SJC); BLiVL Gila Lower Box fishing area, 
along Gila River and rocky- hills above river niwi'in ^.oiranunitt and de,>.it iiS^lmJ 4000 ft, 28 
Apr 2010, Heil & McClain 32322 (SJC); San Simon Valley between Rodeo and Arizona-New 
Mexico line, fine textured soils of the valley bottom in desert grasslands, 4100 ft, with Hilaria mutica, 
Scleropogon brevifolius, Bouteloua eriopoda, Ephedra trifurca, Prosopis, Gutierrezia sarothrae, 10 
April 1978, Moir 1 01 (NMC). Luna Co. : ca. 1 1 mi NW of Florida Station, among rocks in bed of dry 
creek, 29 Apr 1947. McVaugh 8128 (SMU, TEX). 

Collections examined. MEXICO. Chihuahua. Extreme NW corner [of the state], about 50 

m E and 1.5 mi. S of U.S. border, in silty, heavily overgrazed bottom of draw, with Yucca elata, 
Sporobolus airoides, Astragalus wootonii, 13 May 1980, Spellenberg & Ward 5525 (NMC). Sonorsi. 
6.7 mi b\ i j i.l (Mexico li) N ol \Ii Jj1u>« rocky hill- desert skubs 7 *m i'*i3 Eelger 17451 
(ARIZ); vicinity of Cerro Pelon, ca. 5 mi SE of Desemboque, 21 Apr 1968, Felger 17894 (ARIZ); 
near El Guayabo on road 18 km E of Alamos. 27° 0' 20" N, 108° 47' 10" W, 250 m, 16 Mar 1989, 
Ferguson s.n. (ARIZ); along Hwy 89, 9.2 mi S of Arroyo Los Ajos, 7.4 mi N of Mututucachi, 
roadside in oak savanna, 212 Apr 1995, Fishbein 2242 (ARIZ); Rio Mayo, San Bernardo, arroyo, 
Lower Sonoran, margin of a wash, 22 Feb 1935, Gentry 1339 (ARIZ.); Carbo, 50 km N of 
Hermosillo, wash near Mex Hwy 15, 6 Apr 1975, Helmkamp s.n. (UCR); Rancho El Aguilar Noria, N 
of Ures and Santiago, 29° 33' N, 110° 25-26' W, open, broad drainage, Sonoran desert scrub, on mesic 
N slopes, ca. 500 m, common, 21 Apr 1991, Joyal 1995 (TEX); Dto. de Altar, Picu Pass, 23 Mar 
1926, Long 7a (ARIZ); 18.7 mi W of Rte 19, along turnoff 3.2 mi N of Esqueda, oak grassland, 
mesquite bottomlands, 27 Mar 1970, McGill 64048 (ASU); 4.6 mi S of Cucurpe, cliff-face along San 
Miguel River and road, 29 Mar 1970, McGill & Pinkava 6519 (ASU); Near El Guayabo on road 18 
km E of Alamos, 250 m, 16 Mar 1989, Martin & Ferguson s.n. (ARIZ); Mpio. Soyopa, E side of Mex 
16, 1 km S of Rio Yaqui Bridge, 28° 31' 30 " N 109° 32 W, 180 m, 14 Mar 19SS,Martin, Ferguson, 
&Moore s.n. (NMC); NE side of Rio Yaqui bridge on Mex 16, just S of Tonichi, 200 m, in sand, 18 
Feb 1997, Reina G. 97-44 (ARIZ); Arroyo La Quema, near Tepoca, tropical decid. forest on slopes, 
rocky stream canyon, 560 m, 21 Mar 1998, Reina G. 98-378 (ARIZ); Mpio. Benjamin Hill, 3.5 km 
SW of Benjamin Hill on road to Palo Alto, Sonoran desert scrub, 2408 ft. locally very common 
annual on disturbed rocky soil, 1 Jan 2003, Reina G. 2003- 286- A (ASU); 0.8 km N of Mex 16 on 
road to San Antonio de la Huerta, 28° 34' 16" N, 109° 34' 52" W, very open thornscrub, 299 m, 
locally uncommon on flats, 15 Mar 2005, Reina G. 2005-256 (TEX); Agua Prieta, Hwy 2, ca. 26 mi E 
of Agua Prieta and 24 mi W of the state line at Puerto San Luis, desert scrub of Juniperus, Acacia. 
Prosopis, Yucca baccata, Gutierrezia, etc. on rocky volcanic hills, 1300 meters, 19 Mar 1984, 
Sanders 4712 (UCR); Alamos, Rio Mayo Region, Rancho La Huerta, ca. 2 mi NW of Alamos on the 
road to San Bernardo, north of the Alamos airstrip, weedy disturbed areas near buildings and roads, 
420 meters, 15 Mar 1993, Sanders 13152 (UCR); Alamos, Rio Mayo region, roadside c. 12 km W of 
Alamos on the road to Navojoa, in vicinity of Canon Agua Marina, at the foot of the Sierra de 
Alamos, burned roadside in hilly country 1640 ft, 15 Mar 1993, Sanders 13180 (UCR); Alamos, 
Parque Chalaton and along canyon bottom above, SW edge of Alamos in foothills of the Sierra de 
Alamos, tropical deciduous forest and cleared areas, 420-450 meters, 17 Mar 1993, Sanders 13365 



: Taxonomy of Apiastrum, Ammosdinum, and Spermolepis 



(UCR); Dist. Alamos, near Cerros, 4 Mar 1933, Shreve 6167b (ARIZ); 8 mi S of Estacion Llano, 3 
Apr 1935, Shreve 7323 (ARIZ): Palm Canyon, 17 mi SE of Magdalena, in Sierra Babiso, (= Cerro 
Cinta de Plata), stream bed,13 Feb 1977, Van Devender s.n. (ARIZ); 4 mi of El Ocuca on Mex 2, 
21.4 mi E of Altar, annual in wash, 10 Mar 1977. Van Devender s.n. (ARIZ); 17 mi SE of Magdalena 
on road to Cucurpe, Palm Canyon, Cerro Cinta de Plata, 15 May 1979, Van Devender et al. s.n. 
(ARIZ); Alamos, Rio Mayo region; Arroyo Mentidero at the crossing of El Chinal Road, near Rio 
Cuchujaqui, 11.5 km (by air) S of Alamos, tropical deciduous forest, 240 meters, 10 Mar 1993, Van 
Devender 93-97 (ARIZ, UCR); Alamos, Rio Mayo region; La Huerta, l.SkmNNE of Alamos on San 
Bernardo Road, 410 meters, 9 Mar 1993, Van Devender 93-216 (ARIZ, UCR); La Huerta, 1.8 km 
NNE of Alamos on San Bernardo Road, common annual in yard, 410 meters, 9 Mar 1993, Van 
Devender 93-21 6 (ASU); 0.4 mi E of Punto Cirio, Sierra Bacha, Sonoran Desert desert scrub, 40 m, 
24 Mar 1995. Van Devender 95-210 (ARIZ. UCR); El Llano de Curea, foothills thornscrub, locally 
uncommon annual, 514 meters, 19 Mar 2004, Van Devender 2004-161 (ASU); 2.2 km SE of Rancho 
Las Borregas headquarters on road to Nogales, SE tributary of Arroyo Planchas de Plata, sycamore- 
oak canyon, 1187 m, 22 Apr 2004, Van Devender 2004-250A (ARIZ); 7.9 mi N of Esqueda, 11 May 
1948, Wiggins 11 777 (TEX). 

2. Spermolepis echinata (Nutt. ex DC.) A. Heller, Contr. Herb. Frankl. & Marsh. Coll. 1: 73. 1895. 

Leptocaulis echinata Nutt. ex DC, Prodr. 4: 107, 1S30. Avium echinatum (Nutt. ex DC.) 
Benth. & J.D. Hook, ex S. Wats., Bibl. Index N. Amer. Bot, 412. 1878. TYPE: USA. 
Arkansas. "In Amer. bor. ad Red River" [protologue], T. Nuttall s.n. (holotype: BM digital 
image!; isotype: PH digital image!). 

De Candolle noted ("v.s. ") that he had seen the Nuttall collection. 

Stems 5^10 cm. Leaves: blades broadly ovate in outline, 0.7-2.5 cm, 3-pinnately compound, 
ultimate divisions filiform, 2-18 mm x 0.5-1 mm; petioles 3-20 mm. Peduncles (l-)2-5(-6.5) cm. 
Umbels axillary mostly at distal nodes, al! pedunculate; invoiucel bractiets l-3(-4), linear. 1-3 mm, 
margins scabrous-toothed; fruiting rays 5-9(-12), (0)1-15 mm (central umbellet sessile to subsessile), 
unequal, suberect and evidently clustered; umbellets (l-)3-9-flowered; fruiting pedicels l-6(-7) mm 
(central flowers short-pedicellate). Schizocarps 1.5-2 mm, densely echinate-bristly. 2n = 20 
(Constance et al. 1976; Prairie Co., Arkansas, Demaree 61921, duplicate SMU!). Figure 8. 

Flowering (Mar-)Apr-May(-Jun). Sand, gravel, silt, sandy clay, sandy roadsides and flats, 
disturbed areas, ditches, disturbed sites, pastures, rocky slopes, shell banks, sandstone outcrops, 
beaches, creek bottoms, lake shores, prairies, post oak woods, live oak woods, oak-inesquite 
woodland, desert shrub; (0-)100-300(-1500) m; Ala., Ark, Fla., Ga., 111., Iowa, Kans., Ky., La, 
Miss, Mo, NY, N.C., Okla, S.C, Term., Tex, Va.; Mexico (Coahuila, Tamaulipas). 

Specimen examined. MEXICO. Coahuila. Piedras Negras, Pringle 8309 (fide Villarreal 
2001, voucher not seen in present study). Tamaulipas. 20 mi W of Reynosa, desert scrub in clayish 

soil, 28 Feb 1944, Painter & Barkley 14378 (TEX). 

Attributions of Spermolepis echinata to Arizona, California, and New Mexico have been 
based on collections identified here as S. lateriflora. In Texas, typical S. echinata reaches as far west 
as Brewster, Culberson, Jeff Davis, Pecos, and Presidio counties, but it does not extend to El Paso Co. 
at the easternmost extension of the distribution of S. lateriflora. No confirmed records of S. echinata 
exist from New Mexico. 

A plant collected in north-central Texas has the habit and inflorescence of Spermolepis 
echinata and echinata-like fruits (densely tuberculate-hairy) but with the hairs relatively short and 
without an uncinate apex. Wilbarger Co.: 14.5 mi W of Electra, Waggoner pastures, turn W 0.6 mi S 
on Hwy 85, tall grass in draw, mesquite savanna, sandy loam, 12 May 1945, Whitehouse 9828 (SMU). 



: Taxonomy of Apiastrum, Ammosdinum, and Spermolepis 



3. Spermolepis hawaiiensis H. Wolff, Repert. Spec. Nov. Regni Veg. 17: 440. 1921. TYPE: USA. 

Hawaii. Kauai, Weimea, [no date], Hillebrand s.n. (holotype: probably B, Wolff material at 
B mostly extant fide HUH online database). EPITYPE (designated here): USA. Hawaii. 
(Kauai Island): Koai'e Canyon, just below "the fingers" near the ridge and above 
N-facing cliffs W of Lonomea Camp and Kawai'iki and E of Hipalau, 704 meters, 21 Apr 
2004, N. Tangalin 47 (PTBG 043006, digital image on JSTOR!; Fig. 9). This collection was 
made near the type locality. 

Stems 5-20 cm. Leaves: blades oblong to ovate in outline, 1-4 cm, 3-pinnately compound, 
becoming sessile, smaller, and less divided distally, ultimate divisions linear to linear-lanceolate, 3-6 
mm; petioles 10-30 mm. Peduncles 1-3 cm. Umbels at distal nodes, axillary, all pedunculate; 
involucre bracts absent; involucel bractlets (0-)l-5, linear-lanceolate. 1-6 mm; fruiting rays 2-7, (0- 
)5-15 mm (central umbellets sessile to short-pedicellate), unequal, spreading-ascending; umbellets 2- 
8-flowered; fruiting pedicels (0-)2-6 mm (central flower sessile to short-pedicellate), unequal, 
spreading-ascending to ascending. Schizocarps 3-4 mm, densely echinate-bristh', hairs arising from 
multicellular tuberculate bases. 2« = 22 (Wagner et al. 2005). Figure 9. 

Flowering (Dec, Feb-)Mar-Apr. Steq3 mesic forests, gulch slopes and ridge tops in dry 
forest, shrub lands, steep to vertical cliffs, cliffs bases, ridges in coastal dry cliff vegetation, N-facing 
slopes, ridges on bare rock, open, rocky, goat-ravaged area, a'a lava; 50-700 m; endemic to the 
Hawaiian Islands — Hawaii, Kauai. Lanai, Maui, Molokai, Oahu. Information from NTBG (2012) 
and USFWS (2010). 

4. Spermolepis infernensis G.L. Nesom, sp. nov. TYPE: USA. California. San Diego Co.: Hell 

Hole Canyon near Borego, 5-7 Apr 1932, C. Epling & W. Robison s.n. (holotype: RSA!; 
isotype: UC! digital image!). 

Similar to Spermolepis lateriflora in its epedunculate (sessile) umbels but different in its 
sparse fruit vestiture of apically straight, blunt-tipped hairs. 

Stems 7-13 cm, branching from the base. Leaves: blades broadly ovate in outline, mostly 1- 
2 cm, finely ternately dissected, ultimate divisions linear to oblong, mostly 2-6 mm; petioles 10-20 
mm. Peduncles usually absent, occasionally present and 2-3.5 cm. Umbels axillary, sessile at distal 
nodes, sometimes pedunculate at proximal nodes; involucre bracts absent; involucel bractlets (l-)2-4, 
linear, 1^1 mm, margins entire, herbaceous, without scarious margins; fruiting rays 4-5 per node, (0- 
)3-10 mm (inner umbellets sessile or subsessile to short-pedicellate), unequal, spreading; umbellets 
(2-)4-7 flowered; fruiting pedicels 1-5.5 mm (inner flowers short-pedicellate). Schizocarps 1.5-2 
mm, ribs rounded, sparsely to moderately tuberculate to hispidulous-spinulose on the angles and 
intervals with pustulate multicellular mounds, each pustular mound with a straight, erect, blunt-tipped, 
unicellular, hairlike cell, 0.1-0.2 mm or some mounds without a hair, lateral ribs not strongly 
differentiated; oil tubes 3 per interval, barely evident between the thickened ribs. Chromosome 
number not reported. Map 5. Figures 10, 11. 

Flowering Mar-Apr. Desert shrubland; 600-700 m; California (San Diego Co.). 

In deriving the concept and description of Spermolepis infernensis, I have seen only the two 
sheets of the type collection, which include 9 separate plants of consistent morphology. The type 
collection is from the area of the Hellhole Canyon Preserve, which is northeast of Escondido at 
elevations of about 1800-2000 feet elevation. The reference by Epling and Robison to "Borego" 
apparently meant Borrego, since Hellhole Canyon is in the general vicinity of the Borrego Valley and 
the Anza-Borrego Desert State Wilderness Park (see comments above in connection with S. 
lateriflora). 



: Taxonomy of Apiastrum, Ammosdinum, and Spermolepis 



Spermolepis infernensis and S. lateriflora perhaps are sister species, in view of their 
similarity in inflorescence structure and distribution in the western USA It is possible that S. 
infernensis is a recent derivative of S. lateriflora, with a reduction in the density of the fruit vestiture 
and a developmental change that results in truncation of the hairs. The difference in appearance is 
striking, however, and the effect on dispersal potential surely must be significant. 

5. Spermolepis inermis (Nutt. ex DC.) Mathias & Constance, Bull. Torrey Bot. Club 68: 124. 1941. 

Leptocaulis inermis Nutt. ex DC, Coll. Mem. 5: 39, plate 10, fig. B. 1829. Spermolepis 
patens var. inermis (Nutt. ex DC.) Mathias, Brittonia 2: 243. 1936. TYPE: USA. Arkansas. 
"In Amer. bor. ad Red River" [protologue], T. Nuttall s.n. (holohpt BM digital innge'. 
isotype: PH digital image!). 

De Candolle noted ("v. s.") that he had seen the Nuttall collection. 
Leptocaulis patens Nutt. ex DC, Prodr. 4: 107. 1830. Apium patens (Nutt. ex DC.) S. Wats, Bibl. 
Index N. Amer. Bot, 413. 1878. Apiastrum patens (Nutt. ex DC.) Coulter & Rose, Rev. N. 
Amer. Umbell, 110. 1888. Spermolepis patens (Nutt. ex DC.) B.L. Robins, Rhodora 10: 34. 
1908. TYPE: USA. Arkansas. "In Amer. bor. ad Red River" [protologue], T. Nuttall s.n. 
(holotype: BM digital image!; isotypes: NY digital image!, PH digital image!). 

De Candolle noted ("v.s. ") thai he had seen the Nuttall collection. 

Stems 8-80 cm. Leaves: blades oblong-ovate in outline, 3-5 cm, 3-pinnately compound, 
ultimate divisions filiform, 3-30 mm x 0.1-1 mm; petioles 4-15 mm. Peduncles 2-7 cm. Umbels 
terminal and axillary, all pedunculate; involucre bracts absent; involucel bractlets 1-4, linear to 
linear-lanceolate, 2-5 mm, margins scabrous-toothed; fruiting rays 5-11, suberect and evidently 
clustered, unequal, 1-13 mm (central umbellet subsessile); umbellets 2-7-floweied; fruiting pedicels 
(0-)l-6 mm (central flower sessile to short-pedicellate). Schizocarps 1.2-2 mm, tuberculate, 
without trichomes; oil tubes 1 per dorsal interval. Chromosome number not known (reported in 
error as In = 22 by Bell & Constance 1957; voucher from Florida identified here as S. divaricata). 
Map 3. Figure 13. 

Flowering Apr-May (-Jun). Sand, river silt gravelly soil, clay loam, sand prairies, blackland 
prairies, shale glades and barrens, rocky ridges, granite outcrops, limestone crevices, oak-juniper 
woodland, ditch banks, woods edges, roadsides, fields; 30-200(-900) m; Ala, Ark, 111, Ind, Iowa, 
Kans, La, Md, Minn, Miss, Mo, Nebr, N.Mex, Okla, Term, Tex.; Mexico (Coahuila). 

East of the Mississippi River in the southeastern USA Spermolepis inermis occurs only in 
widely disjunct localities in Mississippi, Alabama, and Tennessee; in Louisiana and Alabama, the 
scattered populations occur mostly in prairie patches. Bonafide records seen in the present study are 
documented here. Mississippi. Monroe_Coj ca. 3 mi SSW of Prairie along Hwy 25, 2.4 mi S of jet 
with Hwy 382, 7 May 1996, McDonald 9364 (VDB); ca. 1.5 mi SW of Aberdeen, remnant prairie 
roadside at jet state hwys 25 and 382, 23 May 1996, McDonald 9466 (BAYLU). Oktibbeha Co. : 
Botanic Garden of the South, ca. 1.5 mi S of Sessums on Sessums Rd, Black Prairie region, 21 May 
1997, Leidolf 1512 (VDB). Alabama. Sumter Co. : Blackland prairie between Gainesville and 
Alabama 17, 20 May 1975. Krai 55614 (VDB), Tennessee. Warren Co.: 1 mi NE of Morrison, 
ballast of railroad adjacent swamp and Hwy 55, 18 May 1987, Patrick & Wofford 2029 (SMU). 

Spermolepis inermis probably is not native in Maryland, where it has been reported (Brown 
& Brown 1984). Radford et al. (1968) noted its occurrence in New Hanover Co, North Carolina, but 
no voucher exists in the NCU herbarium (Alan Wealdey, pers. comm.). Some Alabama collections at 
VDB previously identified as S. inermis are instead S. divaricata (Baldwin Co, belong 7712: Dallas 
Co, Sessler 1 1 78; Geneva Co, Krai 90807), and this mistaken identity probably also has been the 
case for the Maryland and North Carolina reports. 



: Taxonomy of Apiastrum, Ammosdinum, and Spermolepis 



Additional specimens examined (outlying populations). USA. New Mexico. Eddy Co.: 
Carlsbad Caverns Natl. Park, 1.8 mi WSW of E boundary via sewage lagoon road, 0.7 mi E of 
sewage lagoons, small arro}~o lined with shrubs, on bajada escarpment, 3650 ft, 19 Apr 1977, Burgess 
4496 (TEX); Carlsbad Caverns Natl. Park, Walnut Canyon, ca, 0.2 mi S, 0.2 mi W of BM 3901, 
gravel alluvium dominated by Brickellia herniate:, 17 May 1977, Burgess 4556 (ARIZ, TEX). 
MEXICO. Coahuila. Rio Grande, Tule Canyon, on Coahuila side above Upper Madison Falls, 
calcareous gravelly soil, Dasylirion, Yucca. Molina, Rhus, Acacia, 475 m. 10 Apr 1973, Johnston et al. 
10615 (LL); San Rosendo Canyon (flows into Rio Grande opposite Brewster Co.), 500-700 m, 
calcareous gravelly loam, Dasylirion, Yucca, Molina, Larrea, Acacia, Quercus, Prosopis, 9 Apr 1973, 
Johnston et al 10597B (LL); Musquiz, spring 1935, Marsh 113 (TEX); Sierra de Santa Rosa. Canon 
El Puerto, Rancho El Puerto, 28 24 N, 101 54 W, matorral de Acacia coulteri, Pithecellobium pollens, 
Zanthoxylum fagara, Yucca thompsoniana, y Opuntia lindheimeri, 900-1000 m, 6 Jun 1991, 
Villarreal 5963 (BRIT): Mpio. Musquiz, Hacienda La Rosita, 26 Jun 1936, Wynd & Mueller 295 
(ARIZ). 

Fruits of a collection from south-central Texas, identifiable as Spermolepis inermis in every 
other way, have some tubercles producing very short, blunt-tipped hairs, similar to those of S. 
organensis and S. infernensis. Wilson Co. : 8 mi S of Elmendorf, 4 May 1945, Cory 48795 (SMU). 

Spermolepis inermis, S. divaricata, S. diffusa, S. laevis, and S. organensis appear to be 
closely related among themselves. If the ancestral species of Spermolepis had hairy fruits with (e.g., 

5. echinata, S. gigantea), then S. inermis can be understood as derived through loss of the hairs. 
Spermolepis laevis probably is a derivative of 5. inermis, through further loss of the surface 
ornamentation of the fruits. Spermolepis divaricata and S. diffusa perhaps are sister species, through 
reduction of the inflorescence from an ancestor shared with A', inermis (maintaining the potential to 
produce fruit hairs). Spermolepis organensis is possibly a peripheral isolate of S. inermis; its short, 
stubby fruit hairs are like those of 5. infernensis and rare populational forms of S. inermis, perhaps 
through partial derepression of the hair formation. 

6. Spermolepis laevis G.L. Nesom, sp. nov. TYPE: USA. Texas. Llano Co.: Enchanted Rock, 

granitic sandy soil, 15 May 1933, E. Whitehouse 11292 (holotype: SMU) 

Similar to Spermolepis inermis in general appearance, especially its strictly pedunculate, 
terminal and axillary umbels and its relatively short, suberect and evidently clustered fruiting rays; 
different in its completely smooth fruit surface, with evident ribs but lacking tubercles on the ribs or 
intervals. 

Stems 8^18 cm. Leaves: blades ovate to broadly ovate in outline, 1-4 cm, 3-pinnately 
compound, ultimate divisions filiform, 4-15(-25) mm; petioles 2-15 mm. Peduncles 2-5 cm. 
Umbels terminal and axillary, ail pedunculate; involucre bracts or very rarely 1-2; involucel 
bractlets 1-4, linear to linear-oblong or linear-lanceolate, 1-3C-5) mm, margins scabrous-toothed; 
fruiting rays 3-8, suberect and evidently clustered, unequal, (0-)2-9(-13) mm (central umbellet 
sessile to subsessile); umbellets (l-)3-8-flowered; fruiting pedicels (0-)3-5 mm (central flowers 
sessile to subsessile). Schizocarps 1-1.2 mm, minutely beaked, surface smooth, dorsal ribs 3, oil 
tubes 1 per dorsal interval. Chromosome number not reported. Map 3. Figure 14. 

Flowering Apr-May(-Jun). Granite outcrops, granitic gravel, limestone gravel, sandy fields, 
oak-cedar slopes, live oak savannas; 200-500 m; Okla., Tex. 

Additional collections examined. Oklahoma. Johnston Co. : 10 mi N of Tishomingo, near 
Wapanucka Road jet, grazed prairie along State Hwy 99, sandy granitic soil from nearby granite 
knobs, 29 May 1948 [fruit], Bobbins 3064 (UC). Texas. Bell Co.: nature prairie near Little River, 5 



: Taxonomy of Apiastrum, Ammosdinum, and Spermolepis 



Jun 1930, Wolff 2201 (VDB); near Little River, field, 11 May 1930, Wolff 2102 (BRIT); Temple, 
from around Substation #5, 11 May 1930, Wolff '21 02 (SMU, perhaps duplicate of the BRIT sheet of 
same number). Burnet Co. : Granite 'hills'[?], sandy soil, 29 May 1922, Thorp s.n. (TEX); near Burnet, 
sandy soil, 26 Apr 1931, Whitehouse 11291 (SMI I): Inks Lake State Parle, ca. 1 mi S of Hwy 29, 
granite outcrop, 1 May 1947, Whitehouse 18439 (SMU). Gillespie Co. : ca. 12 mi N of 
Fredericksburg, dry soil on oak-cedar slope, 29 Jun 1957, Cor r ell & Johnston 17269 (LL). H amilton 
Co. : [no other locality data], 12 Jun 1941. Thorp s.n. (TEX). Llano Co. : [no other locality data], 11 
Jun 1930, Thorp s.n. (TEX). Mason Co. : Mason Mountain Wildlife Management Area, in Middle 
Pasture, 0.2 mi N of Mile-O-More Lake, 23 Apr 2001, Sanchez 2355 (BAYLU); MMWMA, in 
Middle Pasture, near the Lodge, near Una Branta Lake, 19 Apr 2003, Sanchez 3224 (BAYLU); 
MMWMA m West Pasture, downstream from Comanche Lake, near the Beaver Dam, sandy soil, 15 
Apr 2005, Sanchez 3697 (BAYLU); Mason Mountain Wildlife Management Area, in Middle Pasture, 
0.9 mi NE of gate into Headquarters Pasture, sandy soil, 29 May 2005, Sanchez 3879 (BAYLU, 
BRIT); MMWMA in Middle Pasture, 0.3 mi NE of Headquarters Bids., live oak savanna, sandy soil, 
17 May 2008, Hansen 5922 (BAYLU, TEX); granite outcrops on S side of RM 1222, 2.6 km E from 
intersection of US Hwy 87 and RM 1222 at Camp Air, frequent in dry crevice, 17 May 1979, Walters 
301 (SMU). Tarrant Co. : near Fort Worth, in old field, 15 Jul 1924, Ruth 1107 (SMU); gravel road N 
of Crowley under Santa Fe bridge, limestone gravel soil, Grand Prairie. 21 Apr 1946, Whitehouse 
15491 (SMU). 

The Oklahoma collection (Johnston Co.) consists of 5 plants — 3 are typical Spermolepis 
echinata but 2 are typical S. laevis. the fruits completely glabrous. 

From Bell Co., Texas, Wolff 2201 has perfectly smooth fruits; fruits of Wolff 2102 (Fig. lo) 
are very slightly tuberculate but better identified as Spermolepis laevis than S. inermis; Wolff 657 
(BRIT) from Bell Co., however, is clearly S inermis. Both taxa have been collected in Bell, Burnet, 
Gillespie, and Tarrant counties. Field study toward a better understanding of the distribution of 
Spermolepis laevis and its biology, especially its possible interaction with S. inermis, will be 
interesting and useful. 

7. Spermolepis organensis G.L. Nesom, sp. nov. TYPE: USA. New Mexico. Dona Ana Co.: Organ 
Mts., Rock Springs Canyon, NWNW Sec 34, T22S, R4E, common on gravelly loamy granitic 
soil on 5 deg N-facing slope, with Quercus arizonicus, Juniperus deppeana, Garrya wrightii, 
Cercocarpus montanus, Rhus trilobata, 5400 ft, 6 Jun 1995, L Mcintosh 3106 (holotype: 

NMC!). 

Similar to Spermolepis inermis in its strictly pedunculate, terminal and axillary umbels; 
different in its shorter fruiting peduncles and in its corky fruit surface with vaguely formed tubercles, 
some tubercles producing short, straight hairs, some without hairs. 

Stems ca. 20 cm, purple. Leaves: blades broadly ovate in outline, 1.5-2 cm, 2-3 ternately 
compound, ultimate divisions filiform, 4—11 x 0. 1 mm; petioles 2-8 mm. Peduncles 0.9-3.5 cm. 
Umbels terminal and axillary, all pedunculate, indeterminate; involucre bracts absent; involucel 
bractlets 1-4, margins smooth, 1-3 mm; fruiting rays 4-7, suberect and clustered, unequal, (0-)4-10 
mm (central umbellet sessile); umbellets (3-)4-6-flowered, fruiting pedicels (0.2-)l-4 mm (central 
flowers sessile to subsessile). Schizocarps 1.2-1.5 mm, dorsal ribs 3, rounded, often somewhat 
obscured by the corky epidermis, lateral ribs similar to dorsal, oil tubes 1 per dorsal interval; surface 
with vaguely formed multicellular tubercles and' or rounded lateral ridges, tubercles sometimes with a 
straight, erect, blunt-tipped unicellular hairlike cell. Chromosome number not reported. Map 3. 
Figure 12. 



ly of Apiastum, Ammoselinum, and Spermolepis 24 



, 1800 m; N.Mex., known 



Flowering May-Jun. Granitic gravelly loam, oak-juniper s 
only from the type collection. 

The distinctive features of Spermolepis organensis might be interpreted as derived from those 
of S. inermis. The peripheral geographic location of S. organensis (Map 3) also suggests that this 
might be true but other outlying populations of S. inermis exist. Recognition of 5. organensis as a 
species, albeit it weakly defined, emphasizes the apparent, at least partial degression of the fruit liair 
formation, the corky fruit surfaces, and the relatively short fruiting peduncles. 

If S. organensis were interpreted as of hybrid origin from extant parents, the only oilier 
species of Spermolepis currently known from the Organ Mountains is S. lateriflora. The nearest 
known occurrence of 5. inermis is to the east in Eddy Co., that also somewhat of a geographic isolate. 




0# Spermolepis inermis 
Spermolepis organensis 
A Spermolepis laevis 



: Taxonomy of Apiastrum, Ammosdinum, and Spermolepis 



8. Spermolepis divaricata (Walt.) Raf., Bull. Bot., Geneve 1: 217. 1830. Dctucus divaricatus Walt, 
Fl. Carol., 114. 1788. Ammi divarication (Walt.) Pers., Syn. PI. 1: 308. 1805. Aethusa 
divaricata (Walt.) Spreng., PI. Umbell. Prodr., 22. 1813. Sison divaricaium (Walt.) Spreng., 
Sp. Umbell., 113. 1818? Leptocaulis divaricatus (Walt.) DC, Coll. Mem. 5: 39. 1829. 

Babiron divaricaium (Walt.) Raf., New Fl. 4: 24. 1836. Apium divaricaium (Walt.) A.W. 

Wood, Amer. Bot. Fl., 140. 1870, NEOTYPE (Ward 2008, p. 475): USA. South Carolina. 

Charleston Co.: Wadmalaw Island, S of Charleston, 27 May 1988, D. Boufford & E. Wood 

23862 (GH; isoneotypes: MO, NY). 

Walter did not cite a specimen or locality; Ward (2008) noted that "Spm. 40-C, a wispy 

fragment., was labeled "Daucus" by Fraser, and annotated as "divaricatus Walt." by A. Gray, though 

one wonders what he saw that was recognizable." 
Sison pusillum Michx., Fl. Bor.-Amer. 1: 168. 1803. Ligusticum pusillum (Mchx.) Pers., Syn. PI. 1: 

315. 1805. TYPE: USA. [protologue] "In sabulosis aridis Carolinae," A Michauxl (holotype: 

P?). 
Babiron dichotomum Raf, New Fl. 4: 24. 1838. TYPE: USA. Florida. Rafmesque did not cite a 

specimen, noting only "Florida," As synonym of S. divaricata fide Mathias and Constance 

(1944). 
Babiron pusillum Raf, New Fl. 4: 23. 1838. TYPE: USA. Alabama or Georgia. Rafmesque noted 

"sent me from Alabama, and by Dr. Torrey from Georgia as the Daucus pusillus\ see 788." 

As synonym of S. divaricata fide Mathias and Constance (1944), 

Stems 7-40 cm. Leaves: blades oblong to oblong-ovate in outline, 0.5-5 cm, 3-pinnately 
compound, ultimate divisions linear, 3— 10(— 15) x 0.2-1 mm; petioles 1-30 mm. Peduncles 17-40(- 
50) mm. Umbels terminal and axillary, all pedunculate; iff olucre bracts absent; involucel bractlets 
1-3, narrowly lanceolate, 0.5-1 mm, the margins usually callous-toothed; rays 3-6, divaricately 
spreading, unequal, 5-17 mm; umbellets (3-)4-6-flowered; fruiting pedicels (0-)2-9 mm (central 1-2 
flowers subsessile to sessile). Schizocarps 1.5-2 mm, scabrous with minute, upcurved hairs not 
arising from a tuberculate base. Chromosome number: see comments below. Map 4. Figure 15. 

Flowering Mar-Apr(-May). Sandy woodlands (longleaf pine-turkey oak, pine-oak, oak 
scrub, flatwoods, evergreen scub vA ) pia-tK iannant-, sandy peat, sandy roadsides, fields, pastures, 
and clearings, lawns, abandoned gardens, orange groves, moist ditches, swamp and salt marsh edges, 
shell mounds, sand ridges, sandhills. *atsd prairies, sandy peat of bogs; 0-200 m; Ala., Fla., Ga., La., 
Mss., N.J., N.C., S.C., Tex., Va. 

Confusion exists regarding chromosome numbers of Spermolepis divaricata, S. inermis, and 
X echinata. Numbers of 2n = 22 and 2n = 16 apparently both are based on vouchers both identified 
as S. divaricata. Vouchers for both counts were collected in Florida, so at least it is clear that neither 
could have been the basis of a count for S. inermis. The count of In = 20 for S. echinata w r as made 
from an Arkansas plant (see description of S. echinata) securely identified as that species. While it 
seems unlikely that S. divaricata has two such distinct dysploid numbers, the possibility opens an 
interesting evolutionary study. 

In = 22 (Bell & Constance 1957; Florida. Okaloosa Co.: roadside banks just E of bridge 3.6 mi W of 
Crestview, 17 Apr 1954, Bell 1470, NCU [fide A Weakley], VDB!, "Voucher for chromosome count 
of n = 11" on specimen). Initially identified by Bell as Spermolepis divaricata; annotated as S. 
inermis by Mathias & Constance and reported in publication as S. inermis with n = 11; later annotated 
by H.E. Ahles and by Alan Weakley as S. divaricata; confirmed by Weakley (pers. comm.) as S. 
divaricata. 

2» = 16 (Bell & Constance 1957; Florida. Escambia Co.: sandy roadside along Fla. Hwy 297, 5 Apr 
1955, Bell 1514, NCU [fide A. Weakley], "Voucher for chromosome count of n = 8" on specimen). 



: Taxonomy of Apiastrum, Ammosdinum, and Spermolepis 



Initially identified by Bell as Spermolepis echinata and rq>orted in publication as S. echinata with n = 
8 in the Bell and Constance publication. Specimen later annotated by Mathias & Constance and by 

Alan Weakley as S. divaricata; confirmed by Weakley, pers. comm., as S. divaricata. 

Attributions of the species to New Mexico have been based on mis identifications of 

Cyclospermum leptophyllum. PLANTS Database attributes Spermolepis divaricata to New Jersey, 
based on an unpublished "Chrysler Herbarium Checklist" ( by J. Meyer from 

1990. The voucher, correctly identified in the checklist, is this : New Jersey. [Cam den Co. ] : Camden, 
ballast, 26 Jun 1866. C.F. Parker s.n. (CHRB digital image!). The species is regarded here as a waif 
in New Jersey and not a permanent member of the state flora. 

The plants of Spermolepis divaricata from Acadia Parish. Louisiana, from prairie remnants 
along a railroad right-of-way, have pedicels in the upper range of length (mostly 5-9 mm) for the 
species but the central flowers of each umbellet are subsessile and fertile. A cadia Par. : along RR ca. 

3.5 mi SW of Crowley, 7 May 1966, Lemmon 1168 (LSU digital image!). 

Two collections of Spermolepis divaricata are recorded here for Texas. Austin C o. : Industry, 
1895, Mr. H. Wurzlow s.n. (BRIT). Liberty Co. : along Co. Rd 2252 W of the Davis Hill Baptist 
Church and N of Hwy 105 E of Cleveland, 26 Apr 1997, Brown 20285 [without fully mature fruits 
but the pedicels are short and the umbellets have sessile central flowers with evidently maturing fruits] 
(NLU). 

The difference between Spermolepis divaricata and S. inermis can be subtle but is 
nevertheless real. Before full fruit maturation, outgrowths of the ovary surface of S. divaricata can 
look like developing tubercles of S. inermis although they usually have an antrorse orientation. This 
similarity perhaps was the basis for the Mathias & Constance annotation of Bell 1470 (as S. inermis; 
see comments above about chromosome numbers), which lias short hairs and some tuberculate bases 
without hairs. In S. inermis. the central rays are consistently very short and there is a relatively small 
angle of divergence, overall giving the umbels a congested appearance. In S. divaricata, the rays 
(including the central) mostly ars equal to subequal in length and diverge at a relatively greater angle, 
giving the umbels a more open appearance. 

Spermolepis divaricata and S. diffusa characteristically produce a determinate inflorescence - 
- the axillary bud and terminal leaf are suppressed at the distalmost node (Fig. 2B). In some plants 
from Florida, however, this apparent specialization is not expressed (de-repressed?) and the pattern is 
determinate (Fig. 2A). These apparently are populational variants. Examples: Alac hua C o.: Dunn 
525 (FLAS), Rabat 488 (FLAS, 9 plants. 2 determinate. 9 indeterminate), Scudder 1491 (FLAS). 
Baker Co.: West & Arnold s.n. (FLAS). Hernando Co.: Nee et al. 2798 (FLAS). Highlands Co. , 
Baltzell 1 799 (FLAS, 2 plants, 1 determinate, 1 indeterminate). 

9. Spermolepis diffusa (Nutt. ex DC.) G.L. Nesom, comb. nov. Leptocaulis dijfusus Nutt. ex DC, 
Prodr. 4: 107. 1830. LECTOTYPE (designated here): USA. Arkansas. "In Amer. bor. ad 
Red-River" [protologue], T. Nuttall s.n. (BM 001042SS4 diaiul image! isi lectotypes: BM 
001042883 digital image!, NY digital image!, PH-2 sheets digital images!). 
De Candolle noted ("vs. ") that he had seen the Nuttall collection. 

Stems 15-75 cm. Leaves: blades oblong to oblong-ovate in outline, 0.5-5 cm, 3-pinnately 
compound, ultimate divisions linear, 3-15 x 0.2-1 mm; petioles 1-30 mm. Peduncles 20-50 mm. 
Umbels terminal and axillary, all pedunculate; involucre bracts absent; involucel bractlets 1-3, linear- 
lanceolate, 0.5-1 mm; fruiting rays 2—M-6), divaricately spreading, subequal, 15-33 mm; umbellets 
2-4(-5)-flowered; fruiting pedicels (8-)14-32 mm (all flowers of each umbellet with subequal 



pedicels, none sessile or subsessile). Schizocarps 1.5-2 mm, scabrous with minute, upcurved hairs 
not arising from a tuberculate base. Chromosome number not reported. Map 4. Figure 16. 

Flowering Apr-May(-Jun). Sandy clay, sand, roadsides, fencerows, fields, pastures, dunes 
and sandy hills, thin soil under oak-juniper, sandy soil in longleaf pine, pine, oak-pine, oak-hickory, 
post oak, and post oak-blackjack oak w r oods, lake shores; 50-500 m; Ark, Kans., La., Mo., Okla., 
Tex. 




Aptly named Leptocaulis diffusus has long lain in synonymy of Spermolepis divaricata but 
its morphological distinction from typical S. divaricata is easy to discern. The long pedicels and rays 
of S. diffusa usually provide ID-at-a-glance; if the pedicels are in the shorter part of the range, the 
lack of sessile or subsessile flowers provides a second criterion. The range of S. diffusa is entirely 
west of the Mississippi River. The two species appear to be sympatric in central Louisiana and 
southeastern Texas. Vouchers documenting the occurrence of both species in Natchitoches and 
Vernon parishes (Louisiana) are atNLU. 



Plants in a few scattered Texas collections have abnormally short pedicels: Anderson Co., 
pedicels 8-11 mm, Bridges & Kindscher 13731 (BRIT, TEX):. Comanche Co., pedicels 6-10 mm. 
Skinners 20078 (SMU). In the Anderson Co. collection, a low percentage of the umbellets produced 
a single central, subsessile flower that did nor develop a mature fruit. These plants are within the 
geographic range of S. diffusa and are regarded here as populational variants of that species, perhaps 
reflecting ancestal (S. divaricata-ake) characteristics. 



: Taxonomy of Apiastrum, Ammosdinum, and Spermolepis 



The doss similarity and probable sister relationship of Spermolepis diffusa to S. divaricata 
might be emphasized in treating the two as eonspecific varieties. The course here emphasizes their 
distinction in morphology and geography (and by inference, ecology). Pointed field observations in 
their region of sympatry would be interesting, and a chromosome count for S. diffusa might provide 
evidence for an internal isolating mechanism, especially in view of the apparent lability in 
chromosome number within the genus. 

10. Spermolepis castellanosii Perez-Mor., Lilloa 5: 32, fig. 1. 1940. LECTOTYPE (designated here): 

ARGENTINA. Prov. Rio Negro. San Antonio Este, 21 Nov 1928, A. Castellanos (BA 
28/1184). Perez -Moreau cited three other collections — two from Neuquen (leg. Ragonese, 
Perez-Moreau) and one from Mendoza (leg. Ruiz Leal). 

Stems ca. 4-8 cm tall, simple or few-branched mostly at nodes above the base. Leaves: 
blades broadly ovate in outline, 25^0 mm, ultimate divisions 4-8 mm, scaberulous on margins and 
nerves; petioles 7-10 mm, searious-margined at base. Peduncles 20-50 mm. Umbels axillary, all 
pedunculate; involucre bracts absent or 1; involucel bracelets 3^1, linear-lanceolate, entire, 1.5-7 mm, 
unequal; fruiting rays 3-5, 1-15 mm (inner 1-2 umbellets short-pedicellate), spreading; umbellets 3- 
5-fiowered; fruiting pedicels (1— )7— 9 mm (inner flower short-pedicellate), loosely convex to irregular. 
Schizocarps oblong-ellipsoid, attenuate toward the apex, 3.2-5.3 mm, hispid-hirsutulous on the 
angles and intervals with narrowly triangular, straight, blunt-tipped hairs, dorsal ribs 3, rounded, 
lateral ribs not expanded; oil tubes 1 (rarely 2-3) per interval, 2 on the commissural face. 2n = 64 
(Constance et al. 1976; Hunziker 12523, Cordoba, Argentina). Figure 17. 

Spermolepis castellanosii apparently is endemic to west-central Argentina. Perez-Moreau 
(1940) cited collections from the provinces of Rio Negro, Neuquen, and Mendoza. The hexaploid 
chromosome count by Constance et al. (1976) was from Prov. Cordoba. Photos on Flickr by Joseph 
Fourier (2009) are from Prov. San Luis. 

11. Spermolepis gigantea (Coulter & Rose) G.L. Nesom, comb. nov. Ammoselinum giganteum 

Coulter & Rose, Contr. U.S. Natl. Herb. 7: 89. 1900. TYPE: USA. Arizona. Maricopa Co.: 
mesas near Phoenix, 17 Jun 1882, C.G. Pringle 28 (holotype: GH; isotypes: fragment JEPS 
digital image!, NY digital image!, US digital image!). 
Ammoselinum occidentale Munz & Johnston, Bull. Torrey Bot. Club 52: 224. 1925. TYPE: USA. 
California. Riverside Co.: "Hayfields" [Hayfield pumping plant locality], Chuckwalla Valley, 
Colorado Desert, locally abundant in heavy soil of a dry basin under shrubs and in the open, 
500 ft, 13 Apr 1922. P.A. Munz & D.D. Keck 4930 (holotype: POM digital image!; isotypes: 
BM digital image!, JEPS!, UC!). 

Stems 8-26 cm, simple or branching from the base. Leaves: blades obovate to broadly ovate 
in outline, 12-25 mm, ultimate divisions linear, (1— )4— 13 mm; petioles 3-25(3-0) mm. Peduncles 2- 
5.5 cm or penultimate umbel sometimes sessile (see NY isotype, Fig. 19. Umbels terminal and 
axillary; fruiting rays 4-10, (0-)2-22 mm (inner umbellet sessile), unequal, spreading; umbellets 1- 
10 flowered; pedicels (0-)2-8 mm (inner flower sessile to subsessile); involucre bracts (0-)l-3, 
linear or sometimes 3-fid; involucel bractlets 1-6, linear to linear-lanceolate, entire or less commonly 
2-3-fid, 2-12 mm, sometimes searious-margined at base. Schizocarps narrowly elliptic-ovate to 
urceolate-oblong or ovoid-oblong, 3-4 mm, ribs low-rounded, hispid-hirsutulous on the ribs and 
intervals with sharp-pointed, 1-2-celled hairs arising from a conical, non-pustulate base, dorsal ribs 3, 
cordlike and thickened, lateral ribs flattened and broad, nearly obscuring the commissural sulca; 
dorsal oil tubes 3 per interval, commissural oil tubes 2. 2n = 38 (from label of McKay 64: "n = 19 
Chromosome vouchers cultivated in University of California Botanical Garden, C-775"). Map 5. 
Figures 18, 19, 20. 



n: Taxonomy of Apiastrum, 



Calif. 



Flowering Mar-Apr. 



i, sandy flats, desert shrubland with Larrea; 200-800 m; Ariz., 



ire and represented in Arizona only by the collections cited here 
from Maricopa, Pima, and Pinal counties; in California it is known only from the type of 
Ammoselinum occidentale, collected in eastern Riverside County (Map 4). Wolff (1 927) and Mathias 
and Constance (1945) identified Pringle 8314 from Coahuila as Ammoselinum giganteum, but that 
collection, far-distant from the main range of that species, is identified here as typical Ammoselinum 
popei (see citation above). 

Collections examined. USA. Arizona Maricopa Co. : 39 mi NW of Wmtersburg, sandy flat 
among Larrea, 22 Mar 1936, Wiggins 8431 (UC). Pima Co. : ca. 10 miE of Tucson onNogales Road, 
On open roadside, 2500 ft, 27 Apr 1945, Gould 3064 (ARIZ). Pinal Co. : near Casa Grande, 3 Apr 
1937, Darrow s.n. (ARIZ), Casa Grande, 1400 ft firs faintly Pastinaca-scented, 2 May 1965, McKay 
64 (ARIZ); Eloy, 25 Mar 1 930, Peebles, Harrison, & Kearney 6496 (ARIZ-2 sheets). 

Spermolepis gigantea is characterized by these features: umbels terminal and axillary, all 
pedunculate; involucel bracelets long, s carious -margined on the proximal 1/3-1/2; involucre bracts 
usually present, often ternate; peduncles, rays, and pedicels minutely hispidulous on the angles; and 
fruits relatively long, narrowly elliptic-ovate, hispid with non-pustulate-based hairs. The broad 
lateral ribs are perhaps the reason that is has been treated as a species of Ammoselinum. 




©Spermolepis gigantea 
•Spermolepis infernensis 



Map 5. Distribution of Spermolepis gigantea and S. infernensis. 



: Taxonomy of Apiastrum, Ammosdinum, and Spermolepis 



Munz & Johnston (1925) included Ammosdinum giganteum and A. occidentale as the two 
members of Ammosdinum sect. Hesperosdinum Munz & Johnston, but Mathias and Constance (1944, 
p. 104), without comment, placed A occidentals in the synonymy of A. giganteum. where it has since 
resided. Munz and Johnston observed that Ammosdinum occidentale differed from A giganteum in 
its "lower more compact habit, unbranched stems, smaller more compact umbels, pubescent (rather 
than conspicuously callous-toothed) smaller carpels, and twice as many commissural oil tubes." In 
the observation here, however, the fruit vestiture is the same in both taxa, and Mathias and Constance 
(1944) treated A. occidentale as a synonym of A giganteum. Mathias and Constance described the 
lateral fruit ribs of A giganteum as having "corky appendages," but in the observation here, they are 
elaborated hardly more than the dorsal. 

ACKNOWLEDGEMENTS 

I am grateful for loans (to TEX) from ARIZ, FLAS, NMC, RSA, SJC, SRSC, and UC-JEPS 
and for hospitality at BAYLU, BRIT-SMU-VDB, NLU, and TEX-LL while studying there. Thanks 
to Mirta Arriaga at BA for providing close-up photos of the fruit of Spermolepis castellanosii, to 
Liora Spitz at CURB for providing a digital image of the early collection of Spermolepis divaricata 
from New Jersey, to Sarah Taylor and Jochen Schenk at MACF for digilal images of the McFadden 
collection of Spermolepis lateriflora from Los Angeles County, to John Pruski for a digital image and 
close examination of the fruits of Pr ingle 8314 (MO), to Tim Flynn and David Lorence at NTBGfor 
providing the image of Spermolepis hawaiiensis, to Jon Rebman and Mary Alice Kessler for 
observations on the fruit vestiture of SD collections (confirming their identity as Spermolepis 
lateriflora), to Carol Ann McCormick and Alan Weakley for information on NCU specimens, to Sam 
Kieschnick at BRIT for providing scanned images of several specimens, and to Stephen Downie for 
early comments on the taxonomy. This research has been supported in large part by the Flora of 
North America Association. 

LITERATURE CITED 

Bell, C.R. and L. Constance. 1957. Chromosome numbers in Umbelliferae. Amer. J. Bot. 44: 565- 

572. 
Boufford, D.E. 1977. Ammosdinum butleri (Umbelliferae), new to North Carolina. Sida 7: 220. 
Brown, RG. and M.L. Brown. 1984. Herbaceous Plants of Maryland. Port City Press, Baltimore, 

Maryland. 
Bryson, C.E. 1991. Tw ? o weedy species, Ammoselinum butleri (Umbelliferae) and Lepidium 

austrinum (Cruciferae), new to Mississippi. Sida 14: 506-508. 
Calflora. 2012. The Calflora Database. Information on California plants for education, research and 

conservation, based on data contributed by dozens of public and private institutions and 

individuals, including the Consortium of Calif. Herbaria. Berkeley, <http://www.calflora.org/> 
Chodat, R and E. Wilczek. 1902. Contributions a la flore de la Republique Argentine. Bull. Herb. 

Boissier ser. 2. 2: 281-296, 475^190, 521-544. 
Constance, L. 1993. Spermolepis. Pp. 165, in J.C. Hickman (ed.). The Jepson Manual: Higher 

Plants of California. Univ. of California Press, Berkeley. 
Constance, L. and M. Wetherwax. 2012. Spermolepis. Pp. 199, in B.G. Baldwin, D.H. Goldman, 

D.J. Keil, R Patterson, and T.J. Rosatti (eds.). The Jepson Manual: Vascular Plants of 

California (ed. 2). Univ. of California Press, Berkeley. 
Constance, L., T.I. Chuang, and C.R. Bell. 1976. Chromosome numbers in Umbelliferae. V. Amer. 

J. Bot. 63: 608-625. 
Coulter, J.M. and J.N. Rose. 1900. Monograph of the North American LTmbelliferae. Contr. U.S. 

Natl. Herb. 7: 5-256. 
Coulter, J.M. and J.N. Rose. 1909. Supplement to the monograph of North American Umbelliferae. 

Contr. U.S. Natl. Herb. 12: 441-451. 



: Taxonomy of Apiastrum, Ammosdinum, and Spermolepis 



Downie, S.R., K. Spalik, D.S, Katz-Downie, and J. -P. Reduron. 2010. Major ciades within Apiaceae 
subfamily Apioideae as inferred by phylogenetic analysis of nrDNA ITS sequences. PI. 
Divers. Evol. 128: 111-136. 
Fourier, J. 2009. Spermolepis castellanosii. Flickr. Two photos, 1 Nov 2009, from Prov. San Luis, 

Argentina, <http://www.flickr.com/photos/stationalpinejosephfourier/4131897242> 
Gonzalez, A 2010. Fotos de flora nativa de Uruguay y adventicias. Flickr. 

<http://floranativadeuruguay.blogspot.com/2010/12/ammoselinum-rosengurtii-apiaceae.html> 
Keener, B.R. 2007. Noteworthy collections : Alabama. Castanea 72: 47-48. 
Mathias, M.E. and L.Constance. 1944. Apiastrum. Genus 10. North American Flora 28B: 71. 
Mathias, M.E. and L. Constance. 1944. Spermolepis. Genus 11. North American Flora 28B: 71-73. 
Mathias. M.E. and L. Constance. 1944. Ammoselinum. Genus 21. North American Flora 28B: 103— 

104. 
Mathias M.E. and L. Constance. 1950. Four new American Umbelliferae. Bull. Torrey Bot. Club 77: 

133-139. 
Munz, P.A. and I.M. Johnston. 1925. Miscellaneous notes on plants of southern California - IV. 

Bull. Torrey Bot. Club 52: 221-228. 
Munz, P.A (in collaboration with D.D. Keck). 1959. A California Flora. Univ. of California Press, 

Berkeley. 
National Tropical Botanic Garden (NTBG). 2012. Herbarium Database, Herbarium PTBG, National 

Tropical Botanic Garden, Kalaheo, Hawaii, <http://ntbg.org/herbarium/> 
Perez-Moreau, R.A 1940. Una nueva especie de Umbelliferae. Lilloa 5: 31-34. 
Radford, A.E., HA Ahles, and C.R Bell. 1968. Manual of the Vascular Flora of the Carolinas. 

Univ. of North Carolina Press, Chapel Hill. 
SEINET. 2012. Southwest Environmental Information Network Managed at Aizona State Univ., 

Tempe. <http://swbiodiversity.org/seinet/index.php> 
TENN. 2012. Univ. of Tennessee Herbarium website, <http://tenn.bio.utk.edu/vascular/vascular.shtoil> 
Torrey, J.G and A Gray. 1840. Leptocaulis. Fl. N Amer. 1: 608-609. 
USFWS. 2010. Spermolepis hawaiiensis, 5-Year Review, Summary and Evaluation. U.S. Fish and 

Wildlife Service, Pacific Islands Fish and Wildlife Office. Honolulu, Hawaii. 

<http://ecos.fws.gov/speciesProfile/profile/speciesProfile.action?spcode=Q39H> 
Villarreal Q., J. A 2001. Listados floristicos de Mexico. XXIII. Flora de Coahuila. Instituto de 

Biologia, Universidad Nacional Autonoma de Mexico, Mexico, D.F. 

<http://ivww.scribd.com/doc/77962598/Flora-Coahuila> 
Wagner, W.L., D.R Herbst, and D.H. Lorence. 2005. Flora of the Hawaiian Islands website. 

<http://botany.si.edu/pacificislandbiodiversity/hawaiianflora/index.htm> 
Ward, D.B. 2008. Thomas Walter Typification Project, V: Neotypes and epitypes for 63 Walter 

names, of genera D through Z. J. Bot. Res. Inst. Texas 2: 475^186. 
Wolff, H. 1927. Umbelliferae- Apioideae- Ammineae-Carinae, Ammineae Novemjugatae et Genuinae. 

Pp. 1-398 in A Engler (ed). Das Pflanzenreich, Heft 90 (IV. 228). W. Engelmann. Berlin. 



m: Taxonomy of Apiastrum, Ammosdinum, and Spermolepis 3! 




Figure 3. Apiastrum cmgustifolium, representative plant (ASU). 



Nesom: Taxonomy of Apiastrum, Ammoselinum, and Spermolepis 33 



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Figure 5. Ammoselinum rosengurtii (holotype, UC 



Nesom: Taxonomy of Apiastrur, 




Figure 6. Spermoiepis lateriflora, representative collection (AEJ.Z). 



i: Taxonomy of Apiastrum, Ammosdinun, and Spermolepis 36 




Figure 7. Spermolepis lateriflora, representative plant, bottom left of Fig. 5 (ARIZ). 



Nesom: Taxonomy of Apiastrum, Ammoselinum, and Spermolepis 37 




Figure 8. Spermolepis echincita, representative collection (FSU). 



i: Taxonomy of Apiastrur 






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: Taxonomy of Apiastrum, Ammoselinum, and Spermolepis 40 




Figure 11. Spermolepis infernensis, representative plant (from holotype, RSA). 



n: Taxonomy of Apiastrum, Ammose/inum, and Spermo/epis 41 




APIACEAE 
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Figure 12. Spermolepis organensis, holot\pe (NMC). 



i: Taxonomy of Apiastum, 




Figure 13. Spermolepis inermis, representative plant (SMU). 




Figure 14. Spermolepis laevis, representative plant (SMU). 



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Figure 15. Spermolepis divaricata. representative plant (FSU). 



Nesom: Taxonomy of Apiasfrur 



i Spermolepis 45 




Figure 16. Spermolepis diffusa, representative plant (MO). 



i: Taxonomy of Apiastrum, Ammoselinum, and Spermolepis 46" 




Figure 1 7. Spermolepis caste Ikmosii, from the original illustration (Perez -Moreau 1940). 



Nesom: Taxonomy of Apiastrum, Ammoselinum, and Spermolepis 47 




Figure IS. Spermolepis gigctntect, 



m: Taxonomy of Apiastrum, Ammoselinum, and Spermolepis 4§ 




Figure 19. Ammoselinum gigonteum isotype (NY) = Spermolepis gigcmtea. 



i: Taxonomy of Apiastum, Ammoselinum, and Spermolepis 49